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Full text of "Memoirs of the Queensland Museum"

PART 1 (Issued 29 February 2000) 

Ci'D01iI0GHUE,PJ. & ADLARD,.Rl^;v 

Cftt3il<?^e of protozoan parad^ ^^^^^^sMk* i • * i • * j ■ -yl 

BEVEEUDGE.J. ' " ''^ ' ' v-r, r -v *- . * 

•i^w species , ,K--'A- 'v*-»**?@t 

^VEfelQGE,! &CR1BB, T.H. 

NOi&jAmitfiykia (Sci?marda) (PJatyhpiniinthes; PoIycJadida) ftortt Qm^^)^;^^. ^ - < . - 1 i 21^" 

NewZisalaiia , a.32J 

■GLjKp^RD, H.T. & PEMBERTON, M.R. 

Stem casts oi" Leptophio^um aiisiraie (McCoy) Walton an4 the response of their leaf-c^iftio^ 

bases to coffliireHSiEqn- ^ . * . . , ..^j-f . .n-,,.m- .jt ■ .11. . ... ... v-o-;" V • ■ -I ■*L.^7 

COOK,A.G.&BANN,K. ' , v • ^ 

I^ace fos$jl& ^ Upper<CB|$^f@f<hls Jisricftio fK^d^J^tF^ Qu^eMs^ ......... 235 

Atistralikn l^f-tirled geckc^* pftyf^geny, a new genus, twoTiew speetes ari5'bfhe!r''fte^'&te^; 

&)4M|?»,P.J-r. & NG, P.K.L, 

Bountiana. a new genus for Eripfdfl nofrfplcemis Grant & McCulloch, 1907 (Crustacea: 

Brachyura: Criphiidael- * ^ *— v-i.^.i -.,^. >.i - ^ I'fk- At^...,,..^;., 

DAVlES,y.T.& LAMBKIN, C.L. . ^. . . , ' 

^^^mneh aiiew spider genus (Afg^^^if&ta^itP^^^ t}ij& wet1iq|K^r 

G^Kumae of the Mar^haiilialeSrS^ Wlipa ^pi^mtiidli (Ml^^ Erc>mai^ 

Basin, Queensland 285 

& FRITH. D W. 

BoV/er s> stem and stnigtures of the Golden Bowerbird, Friotiotiura fiewtoniam 

PRjm<5.a.&PRlTH,D.W. t S'* ' 

Ji^t^an^is^jetis and behaviour at bowers^qgjf^idal^'Q^I^^ 

satjJi^^i^B..^^ • - 

* ' Jtptne range and associated sociobiology and ecology ofmate Golden Bowerblrds 

Prionodwa newtomam (Ptilonorh> nchidae) 343*. 

Some new (7ivetian (late Middfe Devonian) gas&tipods from ttje Paffrath area ^Ber^i^che^ 

Land, Genna^>^ , . ^. i-^^jn^ ^^^^-h t'aV4^^« f Tt?.V ' >tl''^ ' irW 

JIAR3^a*{STElN, V. & t 

Freshwater dalyelliid flatworm, Gieysitoria ^(^crZw» ^pr J5i3?v4^^ J^t^lOcpeja) 

from southeast Qtj£senaia!ad^:Ai!a1|a!ia- ,,1. i^ii, r,-ji-%»>^Hj.-j--v.— -j ■ ..3^^ 

HERBERT, B. & miMS, B.V. 

-^-.pew species of Streptocephaius (Parastrepiocephalusi (Crustacea; A^OstfflCa:. 

Streptocephalidae) from north Queensland, Australia 3S5 

^NES, B,^.^ HANNAH^ D., VENZ, TV1. & EYRE, T. 

NeVir distribution a^d babi^ dats for the vul^rable ti^'Spppdid. PeiimX'ioMUeita 

^!uge, 1974) ;. * 39T 

HQARE^ B.D, & COOK, A G. 

fleN^zuan and Early Carboruferous Pol;yplaCQphQrafrom Western Australia 395 



KO?|^y|It^^^^^^^^^ tiSfe&Irt&ftfvaiVdsr&firil Atetralfe Mf 

J.N.A., KELLY, M. & KENNEDY, J.A. 

A new Clathria (Porifera: Demospongiae; Microcionidae) frarn the \vesTeni Indian Ocean .... 427 
JEDY, J.A. & HOOPER, LN.A. 

l^ew 1iperi0»9f toetifr re(i»?/a from ngrtUem VaiiUatii (TNDrifera; Dempspongiaet 

•tffli^$i^^;t^^ 4*? 

' 9i&iAmis$^6 '^^s^p^^^^&m^^ i . . : * f.4S3 

^pj^Atl & lOHNSOH Jl#i. 

i'trco Uneaia and P W/fti/i? established as valid species Fieptorhinchus (Percifi^nnes:- 

Haemulidae) rw^-':* ,{..^.^-.<i >f,f.>;.;.i^;-f-ii*^.j.it>t,*^»_-j-4 ^ mll 

Md.OyGilLIN,S.,TOSOLINI. A-M-P. &DRiNNAN,A.N^ - . t v 

Revbion ot an Barty Cretaceous maiqi^(>flQmii*d£n1^'M^1»bf^ 

."SiltlQga^^binae (Acarma: Hal^^ 
* ofthirteen new species . r . ;50S 

HalScaridae from the Great Barrier Reef and Coral Sea: the genera Lohnianneila, 
Scaptogmtthides and Scaptogrmthus (Acarina: RaiacaridaeiliflSinaittatalS^ 

PAJJiRSOH R^-. CATO, D.H., JANETZKI^RA. & WILLIAMS^ S.C. _ .V ^ 
Afeadult Dwarf Minke Whale BaiaeH^^i^ei^fdrif Strata t^oep^de-, :ffiflf^^?^«sfeg 

Island. Qu^paslaad — I . - ^ . .^ . . * . . . - . • ^^^7 

I(AVB7*,RJ. . . . .. ^ 

Taxonomica Araneae |t^'0Ja^^QlMli^^ r^^S^ 

RAVEN, RJ. & BAEHR, B.C. " ' ' ' ' ' ^ ' ♦ " ' , ' ^ * 

Revised status of the genO^^^SMir^Rfi^^ -P^ ^ j-i t ^ i^'tt— 

RQSS,A. 4fc2H^WWAN, W.A. ' ' * - . v 

1S€|Jsl8&i^>.^^^^^ t€i^/. . ».^.^.;;lj^- 

Breeding, feeding and arboreality in Faradelma orientalis: apooriy known* vulnerable 

pygopodid from Queensland liilSh'^: frB 4 ♦ v*^ * 4 U t '3 •> ♦ r • • H ? ^-^ 

MA^ OYQK, iife CROWTHER, M.S. 

S^iB^sessment of northern r^ifssec^ifves of the W^;ec/tfinzts.<Acari^^^^ 

(Oxyopioae: Araneac) , . . .637 

^J^l^-S; & MACKNESS, B.S. 

niaterial of Da^^tfirii^dunfruilU firom the Pliocene Chinchilla Local Fauna of 

CASTRO, p.^ " dcia^teje^faci^Byk^^ ^ 

EAlivteft, R.A. &Sft&A.1[avt. . 

Eastern range extension for M9rethjiC(>^t^9<il£f^ ..... : 

dlFFORD, H.T, & DETTMAN, M.E. 

FossH 255!tft5f%A*!?*?9U»* .Hr'Ti^* — i-^-^— •4^^>t-t-*^-H'r*--f ... * 4-t • '^-H^ s^— a.. . - ..23*4 

4t^tmlonema from the Silurian of ftleil^is&sBas&pNipj* ........ ^ . . 2?1 

SHEAi CfM„ SADLTBIL R.v& iOHNSON, R. * , 

"The scincul WtsrSi^^nia mcpheei Wefts & W^linfeton;T9&4 in Queensland 266 

RfitlDES. M.P. & AMEY, A.P. 

New locality for the endangered sheatbtail bat^ Taphozoua trou^hioni Tate, 1 952 i . • . 2W 



SMALES, L.R. 

Nematodes fi-pm fish from the KeppQl Islands, Ke^pp^l <^©eiidaud 384 
McALPIN, S. 

Addtti(^ to h^Tp&tofmm t^tQ^mrmimih Bitgrtftis? KiJggagi-aphic R^^. * . . . ^ . * * « ^ ? 194 

Queensland wSfc^ ^^-v*^-*.*-**:* w , 55$- 

CHILVERS, B.L. 

Southern Right Whakt £#rf^i?i^^^^ft |[^^ni^iniS^ ifJI3):ftt Mi^^l^ i^f^ 

Queensland ^ ............... ^, . 576 



RllOMM)GKATIimAe(ACA]U^ dREiff BARRIER 

REEF,AjUSTRALTA 



I. BARTSCH 

Bart^cfi, 1.2000 06 30: Rhombognathinae 1 AcatirHaJacaridael frorti itie (.ireat Barrier Reet 
AmusllA J4emoirs of tht OuecnslanJAh^^^ 155-203. Bmban.. IssN 0079-8835. 

Sample fh)m tklal awL^ticbl ^soiiifS 

conmlned one spfndBS ^Acbm^^ sa\A \ 1 ofJlhonttognafi^: Threa qC iheso 12 rhoror 
faogpathlne spoai^^^aORtrUsppnapensis Abe. Rhtmba&mt^fiiffni^^ 
sMUi3W Bartscfa, been recorded pre\' iously fi^dtii was oUtsKieeai^n) Atistt&liiS. the 
nine spades R. fyrtomtm sp. nov., R. delicanttus sp. hov., J?, lafhn'dius sp. nov., /?. levi^M 
■spi nov., R. lon^pe^ sp. nov., R. reticulifer sp. nov., R. seminofatjjs sp. tiov.> R fericulm sp. 
nov. and R. vulu/ipcs $p. nov. are described. A key is given lo ihc cattictn Austiatian iIujiu- 
bognathines. O Eastern A ustralia^ Great Bftrner Reef. Halat^aridac, rhomho^iathinfs, tww 
records, new&pec'm, ifeseripvt^r^^: 

Use SansL'h, Fotschini^!>imfi/iii Senckcnherg. Xofkestr. 31. 22007 ffenfilfiffg, Cerm^\ 
fc-mail BaNschi^'meercsforsdmn^ de): 29 September 1999. 



KhQmbogqatbiliB mites inhsifoit intertitbtJ and 
SKalW^btidal su'fetraia and marine andljrdcle- 

isti waters. Rhombognatliines arc ii!gi\orou>. 
accordingly Ihey are found either on algal fronds 
or on substrata witli epiphy tes. Two rhombognalhine 
]gni^v &hontbognaihy^ and bobactTMs^ strB 
ivi^ly spread in me Pacrfic: 34 species 

of Rhowbo^^nafhus and nine of hchactrus are 
recorded from ihe wc.slern Pacific (Abe, 1998; 
Ratlsch, U^^*^ }- Still the fauna of large areas is 
unknown, especially tiiat of the Great Barrier 
]R^11cS501fciS gap of kjiowiedge, J.C. Otto took 

Parte which latef pfty^ to mMtk 12 ihoitibo- 

jpiathine species, representing t^^o usncmi I sohikims 
— 1 species arid Rhonibogriailins - 1 1 species). 
Three of these species had been recorded 
previously, tlie others are ne\\ to science. 

MATFRIAL AND MHTH()f)S 

Jhe rhombognalhine mites from the Great 
featficr Reef Marine Pmk area were collected and 
sorted by J .C, Otto, Australia Institute of Marine 
Science, TowiisvUle. 

Tht miteS Were cleared in lactic acid mi 
mditnted In g^ennejd^y. Hoiotyit^ ftrp 
oepofdtedi^ftiii^li^mw^ 

*l"owns> ill© (MTW, paf4*ype5 and \ ouchcr 
specimens in the IVfiXj, th^QtieensLaud M useum, 
13nsbane(QM). the Weslem Australian Museum, 
Perth (WAM), the Zoological Inslilule and 
Zoological Museum, Hamburg (ZMBX 9ndtb& 
author's halacarid collection (IB). 

Abbreviations us*fd; AP = anterior dorsal 
{tlate; AE ^ &aX3gti^'&^^ tilaBe; AI* - anal 



plate; ds-i to (is-5 = iixs/t to &S&i, pair of domi 
setae; B = eiifmc*a< httitlb^red I io IV: GA = 

genifnanal plate; GO = genital opening; GT' 
genital plate, OC ^ ocular plate(sK P * palp. P-2 
to P-4 = second to Iburlh palpal segment; pas - 
parambulacral s^Jtae; PD = posterior dorsal plate; 
PE = posterior spheral |»latc(s); pgs - p<?n- 
genhal setae^ nunibereid 1 to 5 from anterior to 
posterior; sgs - subgenital setae Legs numbctcU 
1 to IV, leg segments 1 to h are trochanter, 
basiftniur tclofcmur, genu, tibia and tarsus. 

Drawings were prepared iVtth acaitiera lucida. 

Unless stated otherwise, adjunct and adanal scUie 
are iJiown either in dorsal or ui ventral aspect. 

Length offbe id bstnna h ihat &bnf liieMetiat 

margin of the AD to the end (ifllie invj] Mdves^ 
The length of the PC' includes iIk pair of 
posteriorly prt>iccling cones. The position of a 
seta IS given in a decimal system, w iih reference 
to the l^vigK) of a plate trom the anterior to 
posterior maigin^ the pq&iMon of the legs with 
reference lo the Teilg^ of 'flie idiosoma. The 
length of 'A leg segment is that along the dorsal 
margin. In the seialion formula of the legs, the 
number of pas, solenidia and famuli is excluded. 
Unless stat^ otherwise, the setation jbtitmla of 
ihieielbfeiiTdra -preseots ifae mlkni jbei^ idf'^isal/ 
ventral setae. Measurements in tnicroidetrcs 
unless othenvise stated. 

IQtt Rhamhognathus^ the number of adjunct 
seiti943n and FE, the nutnberof {{orig^tid 
setae atid the setation oil the legs tslditwti t© vary. 
Tach description is supplemented frith notes vm. 
generally unilaicrai, variants; the number al 
eases- are »irparentti«ses. 



166 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1 . Isobactnis ponapensis Abe, 1996; A, idiosoma, dorsal, male; B, idiosoma, ventral, female; C, genitoanal 
plate, male; D, ovipositor, female (genital spines of left half dashed); E, leg 1, ventromedial, female. Scale bar = 
50|jLm. 



SYSTEMATICS 

RHOMBOGNATHINAE Viets, 1927 

Isobactrus Newell, 1947 

TYPE SPECIES (by original designation). Isobactnis 
(Lohmann, \ %^9)=AletessetosiisLohraQnx\, 1889. 

DIAGNOSIS. Dorsal plates AD, OC and PD 
present, sometimes flised. AD with pair of setae. 
OC with (rarely 1 ) setae, 2 gland pores and 0-1 
comeae. PD (or area representing PD) with 1-3 
pairs of setae. Adanal setae absent. Ventral plates 
reduced, epimera I and II rarely fused in the 
median; epimera III with 1-2 setae, epimera IV 
with 1 seta which may insert within the striated 
integument. Genital plate not fiased with anal 
plate. Females with 3(-4) pairs of pgs; males with 
32-98 pgs. Gnathosoma short, generally 
concealed in dorsal aspect. Both pairs of 
maxillary setae on rostrum. Palps 4-segmented. 
P-2 with 1 seta; P-3 without seta; P-4 with 3 
(rarely 4) basal setae. Legs shorter than idiosoma. 
Tibiae I and 11 each with a pair of ventral setae; 
generally 1 seta bipectinate and 1 seta smooth. 



Tarsi I, II and FV with 3 dorsal setae each (one 
species with 4 setae), tarsus III generally with 4 
setae, rarely with 3 or 5. Tarsi lack ventral setae. 
Solenidion on both tarsus I and II dorsolateral in 
position. Tarsi I-IV each with carpite (rod-like 
sclerite) between end of tarsus and central 
sclerite. Central sclerite lacks tine-like process. 
The 2 claws smooth or with tines. 

Isobactrus ponapensis Abe, 1996 
(Fig. 1) 

Isobactrus ponapensis Aht, 1996: 17-24, figs 1-4. 

MATERIAL. 9,^,1 tritonyniph (MTQ), Great Barrier 
Reef, Long Island, Whitsundays, sand at 0.5m, 28 
Febmary 1997; coll. J.C.Otto. 9. 1 tritonyniph (QM 
S50961), collection data as above. ? (IB), collection data 
as above. 

DESCRIPTION. Idiosoma of female 322-335 
long, of male 332. Gland pore on AD im- 
mediately anterior to ds-I (Fig. lA). OC wider 
than long. PD large, marginally foveate, reaching 
between OC. Setae ds-2 within striated in- 
tegument. Setae ds-3 to ds-5 on PD, ds-3 and ds-4 
anterior and level with insertion of leg III, ds-5 



RHONfBOGNATKINAR FROM THE GREAT BARRIER REEF 



167 



posterior to the level of leg IV. Epimera I and 11 
with a seta each. Epimera III and IV separated by 
striated integument (Fig. IB); EIII with ventral 
seta; succeeding seta within margin of ElV. 
Female GO surrounded by plate; anterior pair of 
pgs within striated integument, succeeding 2 
pairs of pgs on and in margin of genital plate, 
respectively. Genital sclerites with 2 pairs of sgs. 
Ovipositor with 10 well-sclerotised genital 
spines (Fig. ID); 2 pairs each anteroapically and 
posteroapically, the latter followed by pair of 
spiniform genital spines. Two pairs of 
anteroapical genital spines large, equal in size, 
each with median process flanked by 2 small 
tines. Posteroapical genital spines in shape 
similar to but slightly smaller than anteroapical 
pairs of genital spines. Male GP with 49 pgs; 
genital sclerites with 4 pairs of sgs (Fig. IC). 
Gnathosoma slightly wider than long. Legs 
shorter than idiosoma. Leg chaetotaxy from 
trochanter to tarsus: legs I and II, 1, 2, 3, 2, 5, 3; 
leg 111, 1, 1, 2, 2, 4, 4; leg IV, 0, 1, 2, 1, 4, 3. 
Ventromedial seta on tibia I (Fig. IE) and II 
bipectinate. Apical pair of fossary setae 
delicately furcate. Carpite between tip of tarsus 
and central sclerite solid. Accessory process on 
claws with single tooth. 

Idiosoma of tritonymph 272-278. Setae ds-2 
and ds-3 within striated integument. 
Arrangement of gland pores as in adults. Small 
genital plate with pair of subgenital setae and pair 
of perigenital setae, another pair of pgs within 
striated integument. Shape and setation of legs 
same as in adults. 

REMARKS. Isobactrus ponapensis was described 
originally on the basis of females, males and 
juveniles from Ponapc, Micronesia (Abe, 1996). 
The individimls from the Great Barrier Reef are 
larger than the adults from Micronesia which 
have an idiosomal length of 255-280, and there 
are small differences in the insertion of the three 
pairs of setae on the PD — in the adults from the 
Great Barrier Reef the interval between ds-3 and 
ds-4 is shorter than between ds-4 and ds-5, in 
those from Micronesia the distance ds-3 to ds-4 
and ds-4 to ds-5 is almost the same. 

The ovipositor of Isobactrus ponapensis bears 
10 genital spines whereas the Northern Atlantic 
species /. selosus (Lohmann, 1889) and /. 
uniscuiaUis (Viets, 1939) have 11 genital spines 
(Bartsch, 1975a). 

DISTRIBUTION. Micronesia, Ponape Island, 
from intertidal coarse coral sand (Abe, 1996), 



and Australia, Great Barrier Reef, from shallow 
water sandy deposits. 

Rhombognathus Trouessart, 1888 

TYPE SPECIES (by original designation). Rhom- 
bognathus notops (Ciosse, 1855) = Pachvgmthus notops 
Gosse, 1855. 

DIAGNOSIS. Dorsal plates AD, OC and PD 
present, sometimes fused. AD with pair of setae. 
OC with 2 setae, 2 gland pores, and 0-2 comeae. 
PD with 1-2 pairs of setae. Adanal setae on anal 
plate. Ventral plates often fused. AE with 3 pairs 
of ventral setae plus 0-6 adjunct setae; PE with 1 
dorsal, 3 ventral and 0-3 adjunct setae. Females 
with 1-45 pairs of pgs; males with 7-25 pairs of, 
generally plumose, pgs. Both pairs of maxillary 
setae on rostrum. Palps 4-segmented. P-2 with 1 
dorsal seta in distal half; P-4 with 3 basal setae; 
apically an often spur-like seta. Legs shorter than 
idiosoma. Tibiae with 2 ventral setae, one or both 
bipectinate. Tarsi I-IV with 3, 3, 3-4, 3 dorsal 
setae, respectively, and ventral setae. Solen- 
idion on both tarsus I and II dorsolateral in 
position. Tarsi with 2 claws. Central sclerite 
between claws lacks tine-like process. Carpite 
(rod-like sclerite) present between end of tarsus 
and central sclerite. 

Rhombognathus cyrtonotus sp. nov. 
(Figs 2, 3) 

ETYMOLOCiY. For tlie ciir\ed (kyrtos, Greek) back 
(notos, Greek), in contrast to the rather tlattened idiosoma 
of the majority of Rliombognathus, 

MATERIAL. HOLOT^TE. 6 (MTQ), Great Barrier 
Reef, 19"20.12'S, 149°02.85'E, Elizabeth Reef, medium 
coarse sand at I Dm, 24 December 1997; coll. J.C. Otto. 
PAR.^TYPES. 2 9,1 tritonymph (MTQ), collection data 
as above. 2 9 (QM S50962)' collection data as above. 2 ? 
(ZMH A96/99), collection data as above. 4 2 (IB), 
collection dam as above. OTHER MATERIAL. 6 (WM1 
99/1439), Great Barrier Reef Lizard Island, Site 'Washing 
Machine', coarse sand and rubble at 7m depth, 14 October 
1998; coll. .I.e. Otto. 

DESCRIPTION. Male. Idiosoma 202-211 long; 
holotype 2 1 1 long, 140 wide. Dorsum raised. AD, 
OC and PD separated (Fig. 2A). Plates unifonnly 
covered with faint reticulum, each mesh 
subdivided. AD 57 long, 75 wide; anterior 
margin broadly arched, posterior margin 
rounded; gland pores in lateral margin; posterior 
line of internal muscle scars at 0.67. OC 65 long, 
48 wide; with 2 comeae, 2 gland pores and a pore 
canaliculus; posterior gland pore by 2-3 times its 
diameter removed from lateral margin of OC. 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2. RhomhognathiLs cyrtonotits sp. no\ .; A. idiosoma, dorsal, male; B, idiosoma. ventral, male; C, posterior 
portion of idiosoma. ventral, male; D. gnatiiosoma, ventral, male; E, tip of tarsus iV, ventral (dorsal setae 
dashed), temaiu; I \ lo^ \. nicJial. leniale; G, leg 11, medial, female; H, leg III, ventromedial, female; I, leg IV, 
ventral, female, asc = anal sclerilc; av = anal valve. Scale bar = 50jxm. 



PD 127 long, 97 wide. Plate evenly and delicately ds-1 1 7 long; succeeding setae 7-8 long. Posterior 

reticulated; posteriorly with pair of narrow ridges seta on OC at 0.60. PD in holotype unilaterally 

but no wide costae. Posterior cones extending Twilli2setae,elsewithpairofs!n^esetae.AdaliaI 

beyond median margin of PD. Pair of gland pores setae on anal valves. 

nearposteriormarginofPD. Analscleritessmall, Ventral plates AE, PE, GP and AP fused to a 

squeezed between lamellar anal valves. Setae ventrd sbi^^ld l^ig. tbis shield 169 long. 



RHOMBOGNATHINAE FROM THE GR^AT BARRIER REEF 



169 




FIG. 3. Rhomhognaihtis cyrtonotus sp. no\M A. idiosoma, lateral, female; B. idiosoma, veniral, female; C, 
oviposilor, female (perigenital sclae and genital spines of left side omitted); D, gnathosoma, lateral, female; E, 
fouilh palpal segment, lateral; F. telofemur II, lateral, female; G, tarsus I, lateral, female (medial setae and claw 
omitted); H, tarsus II, lateral, female (medial setae and claw omitted); I, posterior portion of idiosoma, 
tritdiiymph. gsp = genital spines; pa = papilla. Scale bar = SOfun. 



Areas corresponding to AE and PE lack adjunct 

setae. On PE ventral setae shoncr than dorsal 
seta. GO 27 long, 1 7 wide; not reaching the level 
of insertion of leg IV. Perigenital setae plumose, 
arranged trapezoidally; holoiype with 7 and 9 
86^ kteral to GO and 1 pair of basilar setae, 
close together, posterior to GO (Fig. 2C). 
Spermatopositor 42 long, 45 wide; extending 
beyond anterior p^geni^ setae. 

Gnathosoina 56 long, 48 wide; 1 .2 times longer 
than wide (Fig. 2D). Rostrum 19 long, anteriorly 
naiTow&d; 1 pairs of maxillary setae inserted 



ac^ac^ent; loslral tip with one slender and 1 very 
reduced pair of rostral setae. Tectum slightly 

convex. 

Legs I and IV equal in length; approximately 
0.7 of idiosomal length. Insertion ot leg III at 
0.5 1 , that of leg IV at 0.65. Legs as in female (Fig. 
2F-I). Chaetotaxy of trochanter to tarsus: les L 1, 
2, 3, 3, 5, 3; leg U, 1, 2, 3, 3, 5, 3; leg HI, 1 , C2, 3, 
5, 4; leg IV, 0, 1, 2, 3, 5, 3. Tarsus TV with 2 short, 
pectinate spiniform pas (Fig. 2E ). Tarsus III with 
setiform medial pas and short, spiniform lateral 
pas- 



170 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4. Rhombognathus delicatiilus sp. nov., male; A, idiosoma, dorsal; B, idiosoma, ventral; C, posterior portion 
of idiosoma, ventral; D, gnathosoma, ventral; E, leg I, ventromedial; F, leg II, ventromedial; G, leg III, ventral; 
H, leg IV, ventral, asc = anal sclerite; av ^ anal valve. Scale bar = 50)xm. 



Carpites of tarsi I and II 5 long, those of tarsi III 
and IV 6 long. Each claw with accessory process 
but no further tines. 

Female. Idiosoma 223-247 long. Areas with 
striated integument wider than in males; dorsum 
distinctly raised (Fig. 3A). PD somewhat shorter 



than in male. Ventral plates AE, PE and GP fused 
to a ventral shield; AP separated by wedge of 
striated integument (Fig. 3B). AE and PE lack 
adjunct setae. GO not reaching the level of 
insertion of leg IV. GO surrounded by 5 (rarely 6) 
pairs of pgs; anterior pairs of pgs positioned 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



171 



somewhat anterior to the level of insertion of leg 
IV. Genital sclerites each with 2 sgs. Genital 
acetabula obscured. Extended ovipositor approx- 
imately 55 long; v^ilh pair of small, cone-like 
papillae basally and 5 pairs of sclerotised genital 
spines apically (Fig. 3C); each spine 6-7 long and 
ending with 5 tines. 

Palps of gnathosoma extending beyond tip of 
rostrum (Fig. 3D). P-4 with one wide and 2 
slender setae and a spur-Hke process (Fig. 3E). 
Chelicera 62 long; claw wi th serrate dorsal edge. 

Length :w idth ratio of telofemora: 1.7, 1.8, 1.6, 
1.6. Tibiae I and II slightly shorter than 
telofemora I and II. Tarsi I and II slightly shorter 
than tibiae; tarsi 1 1 1 and IV as long as tibiae 1 1 1 and 
IV, respectively. Telofemora I and II (Fig. 3F) 
each with 2/1 dorsal/ventral setae, telofemora III 
and IV with 2/0 setae. Ventrolateral seta on genu I 
long, stout and bipectinate (Fig. 2F); these setae 
on genua II and III short, seta-like; ventrolateral 
seta on genu IV slightly pectinate and larger than 
seta on genu III. Tibiae I-IV with 2, 1, 1, 2 
bipectinate setae. Tarsus I with papillifonu fam- 
ulus, setiform solenidion and pair of doubled pas 
(Fig. 3G): tarsus II (Fig. 3H) without famulus, 
else similar to tarsus I. Tarsi III and IV resembling 
those of male. 

Tritonymph. Idiosoma 185 long. PD smaller but 
with pair of ridges and reticulation as in adults. 
AE and PE separated. GP and AP fiised; this GA 
with 2 pairs of pgs and 1 pair of minute sgs (Fig. 
31). Number of setae of ietis same as in adults; 
telofemora 1-lV with 2/1, 2?!, 2/0, 2/0 setae. 
Variations. Varieties in characters of adults: 
length of idiosoma, 2: 223-247 (10); length of 
idiosoma, 6 : 202-21 1 (2); number of adjunct setae 
on either side of AE: (19), 1(1); number of 
adjunct setae, PE: 0(19), 1 (1); number of pgs on 
either side of GO, 9:5(1 6), 6 (4); number ofpgs 
plus basilar setae in either half, 6 : 7+1 (3), 9+1 (I ); 
number of setae of leg segments 2 to 5: 



segment 


leg I 


leg 11 


leg III 


leg IV 


2 


2(34) 


2(24) 


1(24) 


1(24) 


3 


2/1(24) 


2/1(24) 


2/0(24) 


2/(K24) 


4 


3(24) 


3(24) 


2(1), 3(23) 


-"*(24) 


5 


5(24) 


5(24) 


5(24) 


5(24) 



REMARKS. Rhomhognathus cyrtonotus is 
characterised by the combination of idiosoma 
wide; PD with I pair of setae; AE, PE and GP 
flised in both males and females; AE and PE 
generally lack adjunct setae; female with 5 pairs 
of pgs; in males pair of basilar setae posterior to 
GO; gnathosoma 1.2 times longer than wide; 
telofemora I-IV with 2/1, 2/1, 2/0^ 2/0 setae, and 



tarsal claw with accessory process but without 
additional tines. 

R. cyrtonotus resembles R. cebints Bartsch, 
1983, a species recorded from the Philippines 
(Bartsch, 1983). Females of R. cyrtonotus are 
distinguished from R, cebuus by the number of 
perigenital setae — 5 pairs in R. cyrtonotus^ 1 1 
pairs of setae in R. cebuus. Male R. cyrtonotus 
have the pair of basilar setae posterior to the GO; 
in R. cebuus these setae are level with the 
posterior edge of the GO. 

At low magnification, Rhombognathus cyr- 
tonotus is separated iVom eastern Australian 
congeners by the combination of: idiosoma wide; 
anal sclerites nanow; gnathosoma short; telo- 
femora I and II each with 2/1 dorsal/ventral setae; 
claws with single tooth. In contrast to the majority 
of Rbombogna/hus species, the posterior gland 
pore on the OC of both adult and juvenile R. 
cyrtonotus is not close to the lateral margin but 
removed by 2-3 times the diameter of that pore. 
Males can be separated from congeners on the 
basis of the position of the basilar setae. 

In contrast to the majority of Rhonibognatbus 
species, tritonymphs of R, cyrtonotus have the 
genital plate fused with the anal plate. 

Rhombognathus delicatulus sp. nov. 
(Figs 4-6) 

ETYMOLOGY. For the delicate {delicatuius, Latin) 
ornamentation of the dorsal plates. 

MATERIAL. HOLOTYPE. 6 (MTQ), Great Ban-ier 
Reef, 19^22.36^S, 149°0L05'E, Club21 Reef, coarse sand 
an(j mbble at 15m. 26 December 1997; coll. J.C. Otto. 
PARATYPES. 9, 6\ 1 trilonyniph (MTQ), collection data 
same as above. 9 and c(QM S50963), collection data 
same as above. 9 (WAM 99/1440), 6 (WAM 99/1441). 
collection data same as above. 9, 6 (ZMH A97/99), 
collection data same as above. 3 9 , 2 d , 2 tritonymphs (IB), 
collection data same as above. OTHER MATERIAL. 9, 
6 (MTQ), Great Barrier Reef, 19°20.12'S. 149°02.85'E, 
Elizabeth Reef, coral rubble at 16-26m, 24 December 
1997; coll. J.C. Otto. (IB), Great Barrier Reef, 
19=20.I2\S, 149°02.85'E, Eli/abe* Reef, coai^e sand and 
rubble at 1 Om, 25 December 1 997; coll. J.C. Otto. 2 9 , 2 d 
(IB), Great Barrier Reef, 18°26.36'S, 146°42.24'E, 
Bramble Reef, coarse sand at 5m, 9 April 1998; coll. J.C. 
Otto. 

DESCRIPTION. Male. Idiosoma 223-260 long; 
holotype 235 long, 137 wide. Dorsal plates 
delicately reticulated (Fig. 4A); each mesh 
subdivided. AD 78 long, 75 wide; anterior 
margin rounded: posterior portion of AD tri- 
angular; posterior scars of muscle strings in an 
almost straight line at 0.60 relative to length of 



172 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 5. Rhombognathus delicatulus sp. nov.; A, tarsus I, lateral, male (medial claw and setae omitted); B, tarsus 
II, lateral, male (medial claw and setae omitted); C, tip of tarsus IV, ventral, male (dorsal setae dashed); D, 
idiosoma, ventral, female; E, ovipositor and two of the genital spines (enlarged), female; F, posterior portion of 
idiosoma, ventral, female; G, gnathosoma, lateral, female; H, tip of tarsus IV, ventral, female (dorsal setae 
omitted); I, idiosoma, ventral, tritonymph. gac = genital acetabula. Scale bar = 50|xm. 



AD. Pair of gland pores in lateral margins at the 
level of insertion of leg 1. OC 67 long, 37 wide; 
with 2 comeae and 2 glands pores in lateral 
margin; pore canaliculus almost halfway between 
gland pores. PD 137 long, 85 wide; evenly 
reticulated; each mesh with 12-18 minute pits. 
Anterior portion of PD rounded; posterior cones 
hardly extending beyond median margin of PD. 
Anal valves prolonged, extending beyond narrow 
anal sclerites. Dorsal idiosomatic setae small; 
ds-1 10 long and hardly longer than setae on OC 



and PD. Posterior seta on OC at 0.44 relative to 
length of OC, that equals level of median edge. 
Single pair of setae on PD at 0.26. Adanal setae 
on anal valves. 

AE, PE, GP, and AP fiised to a ventral shield 
(Fig. 4B); this shield 195 long. Integument of 
ventral plates delicately punctate. Portion 
representing AE with 1 pair of adjunct setae. No 
adjunct setae on PE. GO 27 long, 17 wide. 
Anterior margin of GO almost level with 
insertion of leg IV; interval between posterior 



KUOMBOQN ATIUNAE FROM THE 0R£AT BARRIER RE£F 



edge of GO and end of idiosoma 1 .4 times length 
of GO. Perigeoilal seiae plumose; arranged 
trapezoidally; on either side a line of S setae; 
basils setae adjacent to GO and at O.SO relativcr 
Id leng^ of <X}. SpermaiQposuor 40 Ipi^ 4^ 
wide; extending beyond GO and ahteriot 
pcrigenital setae (Fig. 4C). 

C^9l;bo$oQ]a 67 long, 42 wide; 1^ times lofigcsr 
U^iwtd^ fFig.4D). Rostrum slender. 3() long, U 

wide. Basal pair of maxillaiy setae almost IS 
long; succeeding adjacent pair ot setae distinctly 

Leg I inserted at 0.1 1, leg IV at 0.64. Legs 
shorter tlian idiosoma. Leg ! and IV subequal in 
length, about 0.8 tunes of length of idiosoma. 
Form of teloferaora as lu feuuale. Tibiae C3'l- 
indrical; each tibia sli^rtSy ^ortorthan telof^ur 
of that leg. Leg chaetotaxy, from trochanter to 
tarsus: leg I, 1, 2, 5, 5, 5. 3 : leg H, 1, 2, 5,5, 5,3: 
leg in, 1,1,3, 3. 5, 4; leg IV, 0, 1. 3. 3, 5, ?. 
Telofemora I-IV with 4/lT4/I, 3/0, 3/0 setae. 
Mfentrolateral seta on both genu 1 a^ I\' bi- 
pectinate (Fig. 4E,^ tb^se setae on gienua U and 
rn slender (Fig.4F, G). Tfbial with pair of stout, 
bipectinate setae: tibiae II and IIT each wilh 
.slender, smooth ventrolateral and stout, bi- 
pectmate ventrofiiedial seta. Tibia iV with largt^ 
ventrolateral and shorter vervtromedial seta; boUi 
bipectinate. Dorsal setae on tarsi at low 
magnification plain, under oil immersion 
delicate pluinosity nrcogrusable. Solenidion on 
larsus I 10 long, famulus papillifomi. 1 long, 
positioned halfwav between solenidion and 
dorsal fossary seta iFig. 5A). Solenidion on 
tarsus U. lOiqngCFig. aD). TacSi ladld H ^hwith 
pair of dtnAied poa. TMus III widi setrfaml, 
eupathid' lii^Mf.pas and ^piniform. pectinate 
lateral pas. Medial pas on tarsus IV long and 
plumose: lateral paS spbiifitm ^ llltettseiy 
pectinate (Fig. 5C ). 

Carpites on tarsi I and II 6 long, on tarsi 111 and 
TV 7-S long. AcoessQzy process of claws "with. 

single tooth. 

Feniale. Idiosoma 192-269 long. LX")rsal aspoc: 
same as in male, Setae ds-1 slightly longer than 
followmg setae. Ventral plates AE, PE and GP 
^sed; this ventral shield contiguous with ana] 
plate. GO surrounded by XQ p^ngstntil 
Anterior pair of pgs levd With ventral setae on 
epiincTa IV. Second pair of pgs almost IcatI with 
anterior edge of GO (Fig. 5D ). hiterval between 
pgs-4 and pgs-5 slightly larger than between the 
other pgs. Genit^ sclerltes 63 loo^ cacl^ scierite 



gcrutal spines; cacii genital spine with one largo 
medial tooth and 4-Ji btend teetb (F^ 56). 
(rcnital acctabula very small, positioned im* 
mediately posterioir to theievel of pgs-4 {Fig. if\ 

, GoMlltotirba 1.^1.^ liines longer than wide. 
&dp$8l|3dght: expending beyond end of rostnmL 
CFig.'5G). Basal pair oi rostral setae ahnost as 
long as apical pair of roaxillary setde. ChdtCerA 

70 long. Cheliceral claw short. 

Iclotcmora I-IV 19. 2.1J. 2.0. 1.9 times longer 
than high Me^iial ;vhs on tarsus IV setifoou» 
delicately plumose; latexaL pas spinifonn« 
peclinale (Fig. 511). 

Trtktnympk Idios<>ma 1 82-212 long. PD^tfer 
ihan in adults, t il Jim and lateral portions 
foveate, not reticuiaic . Ventra] plates* AE> add 
OA separate. AE with pair of adjmffit-^tae. No 

adjonci scliic on PL GP .nid AP fused (Fi;j 'Ii 
G A with 2 pairs ofpgs and 1 pair of sgs; 3 pairs of 
minute genital acetabula. Leg chaetotaxy fronl 
trochimler io fareus: legs 1 and 11, 1 . 2. 4. 5. 5, 3; 
leg 111. IJ 5. 4; kg I\^ 0, U 2-3. 3, 5, 3. Telo^ 
femoral to IV wiih3''],3/U3.'Oand2-3A) setae. 

Vai'iations. Amongsl the material examined* 
several individuals, e.g. from. ElizabcUi Keef 
(MTQ), differ fron: t>ij ise from the type locality in 
the following charatieib: giiall^osomal base and 
rostrum short, gitaihosoma I3-I.4 times longer 
than wide (Fig. 6A. B'l, reticulation of PD rather 
prointoejiit; ds- , acom nvitc the Icngtli of the 
SUC^^ecKog setae: ds-3 slightly posterior to 
rpedial cotTicr of OC; bipectinate ventral seta nn 
genu 1 as long as ventral setae of tibia 1 fFig. fiC); 
ventral seta on genu IV slender \^v.\ pecuiat* 
iTig, 6D'i; ventromedial seta or. iibia W sicnJcr. 
Its pectirialion very faint or lacking; tcloleraora 
hatdty longer t]ian tibiae. Most marked is the 

sh-::Tt yniathosomii W'idiiii tlie fhombognathincs. 
such a difference in the lei^gth of the gnathosoina 
iinusisaL Nonis&ekss, the specimens outlined 
arc presently not regarded as belonging, to a 
separate species. 

VWecib of chamder? in aduttSr length of idio* 

soma, '?: l92-26*> ilO); length of idiosoma, i^^: 
223-260 (10): number oi adiunct sei;.ie on either 
side of AE: i 2 ), I (3 3 ), 2 ( 5 ); niiin bcr at adj unct 
setae on P& (4U>; nun^ber o£pg;s on either side 
oCtiO^ $ t S^)$liainb<NfOf^g$plcisbaaiIar setae 



174 



MHMUiRS OF THE QUEENSLAND MUSEUM 






FlG.p.RhombognaTtm deiiemulussg^ nov.from Elizabeth Reeft A, ffwthpsema*V€ntraJ. male; B^gnathosoma, 
latei^l^ ^^t\ C, l^t. mteiAl, fbt&ale: leglV^ medial, female. Scale l)ffir»SOMJii* 



in either haU; S: 7+1 f3), 8+1 (10), 9-M (7); 
number of setae of leg segments 2 to 5: 



Mgment 


icgl 


leg 11 


leg III 


leg IV 


2 




2(4iH 


1(40) 


1(40) 


3 


3/1(1), 


4/1(40) 


3/0(3S>), 


j/a(40j 




4/0(1). 




4/0(1 J 






4/1(38) 








4 


5(40) 








5 


5(40) 


5rn)) 


5(40) 


N4(h 



RI:MARKS. The nK)sl obvious characters of 
Rhomhugna/Jiiis ilclicaiidus are: the utiifomily 
reticujiated PD with a single pair of setae; the 
ventral shield: the narrow anal sclerites surpassed 
by the anal valves, the low number of adjunct 
setae; the slender gnathosoma; the combination 
4/1 , 4/1, 3/0^3/0 setaemtheteteffcm^ia I-l V; and 
the claws with a -single tine. At a ^iip^cial 
glance, R. delie&tulus resembles th^ western 
Pacific R. nep tune 1 1 US BavXsch, 1992. /?. oblongus 
Bartsch, 1989a, R. teurinus Abe, 1996, and the 
westem kvs^asSI^Kp&cmm^^ 1993. 

R. ncpinnel/ns and R. fevn'nas differ from R. 
delicQiulua in the outline of tlie AD, the insertion 
of ai5-3 on the "OC* azi!l tBp shape of the 



gnathosoma. The PD of R. ohlongtts is more 
slender than thai of/?, del icat nl as . R- 
psammophiius lacks the reticulation on the dorsal 
plates, the series of muscle scars is closer to the 
posterior majgrn of the AD, j^d tlie ds-4 ate 
dosei: to the ^terior margin ofthe PDlhan in R, 
deHdisttutus. 

Rhombognathns lathridius sp. nov. 

(Figs 7, 8) 

ETYMOLOGY. Because of the shape of the body and 
legs, this species is believed to live hidden {laihricSos^ 
Greek) in sandy deposits. 

MATERIAL. HOLOTYPE. S (MTQ). Great Banier 
Reet: I8°48.92'S, t46°25.76T, Pandora Reef, St Ucoarae 
sand, Im, 22 Januan' I9<)S: coll J.C. Otlo. P/VRATYPES. 
2<J, 1 trilonvmph (M rQl collcclion daui as ahnive. 
c5(QM S50964). collcclion data as above. 2V (WAM 
99/1442, 1443). colleciion dala Lb. abo\e. r'. (/MH 
A98/99), collection data as above. 4 t , 3 d , 2 aiion>Tnphs 
(tB), coUebtim data as above. 

DFSCRIFTION. XJa/e. Idiosoma 229-254 long, 
holotype 248 long, 145 wide. Dorsal plates very 
^t^ andnlinutely teticulated. AD^ QC andPD 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



175 




FIG. 7. Rhombognathus lathridius sp. nov.; A, idiosoma, dorsal, male; B, idiosoma, ventral, male (adanal setae 
illustrated both in A and B); C, posterior portion of idiosoma, ventral, male; D, gnathosoma, ventral, male; E, 
gnathosoma, dorsal, male; F, gnathosoma, lateral, male; G, tarsus II, lateral, male (medial claw and setae 
omitted); H, tip of tarsus IV, ventral, male (dorsomedial seta omitted); I, tarsus III, lateral, male (dorsomedial 
seta, medial pas and claw omitted); J, tarsus I, lateral, female (medial claw and setae omitted), ads = adanal seta. 
Scale bar = 50|jLm. 



separated (Fig. 7A). AD 82 long, 72 wide; 
anterior margin with small, rounded process; 
posterior margin ovate. Pair of gland pores in 
lateral margins at the level of insertion of leg I. 
OC 67 long, 42 wide; each plate with single 
cornea; two gland pores in lateral margin; pore 
canaliculus between pores. PD 145 long, 80 
wide. Anterior portion of PD triangular. Pair of 
gland pores at base of posterior cones. Dorsal 



setae small; ds-1 approximately 10 long, not 
markedly longer than posterior pairs of setae. 
Setae ds-1 on AD posterior to the level of gland 
pores and at 0.52-0.54. Setae ds-3 at 0.52 relative 
to length of OC. Single pair of setae on PD at 
0.27. Adanal setae 7 long, on tube-like pedestals. 

Ventral plates AE, PE, GP, and AP fused to a 
ventral shield (Fig. 7B). Surface of plate 
delicately punctate. AE with 2-3 pairs of adjunct 



176 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 8. Rhombognathus lathridius sp. nov.; A, idiosoma, ventral, female; B, ovipositor, ventrolateral, female; C, 
leg I, medial, female; D, leg II, medial, female; E, leg III, medial, female; F, leg IV, medial, female; G, tip of 
tarsus IV, ventral, female (dorsal setae omitted); H, idiosoma, ventral, tritonymph. Scale bar = 50|xm. 



setae. PE with 1 dorsal, 3 ventral and 2 marginal setae plumose; 7-8 setae arranged trapezoidally 
adjunct setae. GO 25 long, 15 wide. Perigenital around GO; pair of basilar setae posterior to GO 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



177 



(Fig. 7C). Spermatopositor 37 long, 40 wide, 
extending somewhat beyond anterior perigenital 
setae. 

Gnathosoma slender, 75 long, 46 wide, 1.6 
times longer than wide (Fig. 7D). Rostrum 32 
long, 15 wide, almost parallel-sided. Tectum 
truncate (Fig. 7E). Basal pair of maxillary setae in 
basal half of rostrum. Palps slightly flattened, 
extending beyond tip of rostrum. P-2 with long 
dorsal seta. Cheiicera 71 long. Cheliceral claw 
narrow (Fig. 7F), 1 long, its dorsal margin smooth. 

Legs short, without claws and carpite about 
half as long as the idiosoma. Insertion of legs III 
and IV at 0.47 and 0.64, respectively. Number 
and arrangement of setae on trochanters to tibiae 
as in female. Tarsi I-IV with 3, 3, 4, 3 dorsal setae. 
Tarsi I and n each with pair of doubled pas; as in 
female, famulus on tarsus 1 2 long (Fig. 7J), 
solenidion 6 long. Solenidion on tarsus II 1 long, 
conspicuously wide (Fig. 7G). Lateral pas on 
tarsus III flattened, bipectinate (Fig. 71); medial 
pas setiform. On tarsus IV dorsolateral fossary 
seta plumose; lateral pas flattened, bipectinate; 
medial pas plumose (Fig. 7H). 

Carpite short, on tarsi I and II 4 long; on tarsi III 
and IV 5 and 6 long. Claws short and smooth, 
they lack accessory processes and tines. 

Female. Idiosoma 235-254. In dorsal aspect 
similar to male, though areas of striated in- 
tegument larger. Pair of setae on PD at 0.25-0.28. 
Ventral plates AE, PE, GP, and AP fused to a 
ventral shield (Fig. 8A). GO surrounded by 10 
perigenital setae. Anterior portion of GO not 
extending to the level of insertion of leg IV. Four 
anterior pairs of setae positioned almost 
equidistant. Anterior pairs of pgs 15 anterior to 
GO but not extending beyond the level of 
insertion of leg IV. Genital sclerites with 2 pairs 
of sgs. Ovipositor with 5 pairs of claw-like 
genital spines (Fig. SB); spines 5 long, 3 wide; 
each with 5-6 tines. 

Legs I and II flattened. Telofemora I-IV 1.3, 
1.3, 1.2, and 1.3 times longer than high, 
respectively. Tibiae I-IV slightly shorter than 
telofemora. Leg chaetotaxy: leg I (Fig. 8C), 1, 2, 
7, 3, 6, 3; leg II (Fig. 8D), 1, 3, 7, 3, 6, 3; leg 111 
(Fig. 8E), 1 , 2, 4, 4, 5, 4; leg IV (Fig. 8F), 0, 2, 4, 5, 
5, 3. Basifemora, genua and tibiae dorsally with 
short slightly plumose setae which are less than 
length of each segment, and 0-1 long smooth 
setae which are much longer than length of the 
segment. Ventral seta on genua delicately 
pectinate. Tibiae l-IV with 2, L 1,2 bipectinate 
setae. Ventral setae on tibia I equal in length; on 



tibia IV ventrolateral seta slightly smaller than 
ventromedial one. Slender ventrolateral seta of 
tibiae II and III as long as coarsely bipectinate 
ventromedial seta. Dorsal setae on tarsus IV 
smooth; lateral pas short, flattened, pectinate; 
medial pas slender, lightly plumose (Fig. 8G). 
Tritonymph. Idiosoma 232 long. Shape of AD 
and OC similar to that of adults. PD shorter; ds-4 
inserted at 0.22. Plates AE, PE and GA separated 
(Fig. 8H). AE with 1-2 pairs of adjunct setae; PE 
with I pair of adjunct setae; GA with 2 pairs of 
pgs. Leg chaetotaxy, from trochanter to tarsus: 
legl,l,2,6,3,6,3;legll,l,3,6,3,6,3;leglll,l, 
2, 3, 3, 5, 4; leg IV, 0, 2, 2-3, 4, 5, 3. Arrangement 
of pectinate setae on tibiae same as in adults. 
Tarsi III and IV each \\ith flattened bipectinate 
lateral pas, and slender, setifonn medial pas. 
Variations. Varieties of characters in adults: 
length of idiosoma, 9 : 235-254 (9); length of 
idiosoma, cJ: 229-254 (6); number of adjunct 
setae on either side of AE: I (9), 2 (21), 3 (2); 
number of adjunct setae, PE: 1 (10), 2 (22); 
number of pgs on either side of GO, ? : 4 (1), 5 
(18), 6 (I); number of pgs plus basilar setae in 
either half, 5:7+1 (11), 8+1 (1); number of setae 
of leg segments 2 to 5: 

segment leg I leg U leg III leg IV 

2 2 (28) 3(28) 2(27) 2(27) 

3 5/2(28) 5/2(27). 3/1(26), 3/1(26), 

6/2(1) 4/0(1) 4/1(1) 

4 3(28) 3(27), 2(1) 4(26), 3(1) 5(27) 

5 6(28) 6(28) 5(27) 5(27) 

REMARKS. In the shape of the body, gnatho- 
soma and legs, Rhombognathus lathridius is 
similar to R. caudiculus Bartsch, 1983, R. con- 
junctus Bartsch, 1986, R. intermedius Schulz, 
1933, R. latens Bartsch, 1993 and R. latibulus 
Bartsch, 1993. 

R. la/ens, a species known from Western 
Australia (Bartsch, 1993), can be separated from 
the others on the basis of the two pairs of setae on 
the PD and the enlarged number of perigenital 
setae around the female GO. R. conjunctus, an 
inhabitant of the Mediterranean (Bartsch, 1986), 
has the dorsal plates AD, OC and PD fiised to a 
dorsal shield. R. intermedins^ widely spread in 
sandy deposits in the shores of the Baltic, North 
Sea and northeastern Atlantic (Bartsch & 
Schmidt, 1979), has a short gnathosoma, slender 
telofemora and the adanal setae do not stand on 
small pedestals. R. caudiculus and R. latibulus, 
recorded from the Philippines and Western 
Australia (Bartsch, 1983, 1993), respectively, are 
most similar to R. lathridius. but the former 
species bears a single pair of adj unct setae on both 



178 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 9. Rhomhognathus Icvii^aius sp. nov.. niaio; A, idiosonia, dorsal; B, idiosonia, vcnlra!; C, gnathosoina, 
ventral; D, posterior portion of idiosonia, \enlral; E. gnathosoma, lateral (cheiicera dashed); F, leg I, 
ventromedial; Ci, leg H, medial; H, leg III. medial; 1, leg IV, medial, asc = anal sclcrite. Scale bar = 50p.m. 

AE and PE, and the anterior portion of the PD is setae on the PE, and the anterior portion of the PD 
broadly rounded, whereas R. lathridius has 3-4 is triangular. Further distinguishing characters 
pairs of adjtmct setae onfhe AE, 2 pairs of adjunct are: in the male ofR, caudiculus the pair of basilar 



RHOMBOGNATIIINAE FROM THE GREAT BARRIER REEF 



179 



setae is level with the posterior margin of the GO; 
and the telofeniora I-IV ofR. latibidus bear 4/2, 
4/2, 2/1,2/1 selac. 

All specimens of/?, lathridhts have an elongate 
gnathosoma with a slender rostrum; there is no 
tendency of reduction of the gnathosomal length. 

Tritonymphs of R. lathridius have the GP and 
AP fused, a character shared with R. cyrtonotus 
and R. delicatuhis. 

Rhombognathus levigatus sp. nov. 
(Figs 9,10) 

ETYMOLOGY. From levigare to smooth for the 

almost smooth siirtace of the dorsal plates. 

MATERIAL. HOLOTYPE. 6 (MTQ), Great Earner 
Reef, 18°48.92'S, I46°25.76'E, Pandora Reef, St. 1, coral 
rubble. 0.3m, 22 January 1998; colL J,C. Otto. 
PARATYPES. 29,1 tritommph (MTQ), collection data 
as above. d(QM S50965), collection data as above. ? 
(WAM 99/1444), collection data as above. 9 (ZMH 
A99/99), collection data as above. 9, 6 (IB), collection 
data as above. 

DESCRIPTION. Male, Idiosoma 267-276 long; 
holotype 267 long, 1 73 wide. Surface of plates 
almost smooth; integument of lateral portions of 
AD and PD and medial portions of OC pierced by 
minute pores (Fig. 9A). AD 98 long, 93 wide. 
Anterior margin arched; posterior margin 
broadly rounded. Posterior transverse line of 
muscle scars at 0.69. OC 75 long and 48 wide. 
Lateral margin with 2 gland pores and, halfvvay 
between, a pore canaliculus. PD 140 long, 110 
wide; wider than AD, and its anterior margin in 
the median truncate. Very slightly raised pair of 
oblong areolae with minute pores; posterior 
portion of PD faintly reticulated in the median 
and laterally. Anal sclerites not surpassed by anal 
valves. Setae ds-1 approximately 12 long; 
inserted at 0.44. Setae ds-2 and ds-3 on OC; ds-3 
at 0.57. Single pair of setae on PD at 0.28. Adanal 
setae on anal valves. 

Ventral plates A E, PE and GP fused; GP and AP 
contiguous (Fig. 9B). Ventral shield from 
camerostome to tip of anal cone 2 1 9 long. AE and 
PE each with 1 pair of adjunct setae. GO 40 long, 
23 wide. Perigenital setae plumose, arranged 
trapezoidally, with 1 1 setae in a line and 1 pair of 
basilar setae level with posterior edge of GO (Fig. 
9D). Anterior edge of GO slightly surpassing the 
level of insertion of leg IV. Distance between 
posterior edge of GO and end of anal cone 
equalling length of GO. Genital sclerites with 2 
pairs of sgs. Spermatopositor 60 long, 57 wide; 
extending beyond anterior pgs. 



Gnathosoma short, 70 long, 58 wide, 
length :width ratio 1 .2. Rostrum 25 long, 13 wide, 
triangular, shorter than gnathosomal base (Fig. 
9C). Palps tightly appressed to rostrum. Chel- 
icera 75 long (Fig. 9E). Cheliceral claw serrate. 

Length of legs (claws included) 0.7 times that 
of idiosoma. Relative to length of idiosoma, 
insertion of legs III and IV at 0.55 and 0.72, 
respectively. Telofemora 1-lV each about 1.7 
times longer than high. Tibiae 1 and II somewhat 
shorter than telofemora (Fig. 9F, G); tibiae III and 
IV as long as telofemora (Fig. 9H, I). Tarsi 1 and IF 
as long as these legs, tibiae; tarsi III and IV longer 
than tibiae III and IV, respectively. Leg 
chaetotaxv: lea 1, 1, 2, 7, 5, 5, 3; leg 11, 1, 2, 7, 5, 5, 
3; leg IIU'^l, 2,^4, 3, 5, 4; leg IV, 0, 2, 4, 3-4, 5, 3. 
Telofemora I-IV with 5/2, 5/2, 3/1, 3/1 setae. On 
both genu I and II ventrolateral seta longer than 
ventromedial one; both setae almost plain. Tibiae 
I-IV with 2, 1, 1,2 bipectinate spiniform setae. 
On tibia IV ventromedial spine shorter than 
ventrolateral one. Tarsus III with 4 dorsal selac; 
distance between 2 basal ones equalling half 
height of tarsus. Tarsus 1 with 1- longpapillifomi 
famulus and 8- long setifonn solcnidion (Fig. 
lOA). As in female, solcnidion on tarsus II 10-11 
long (Fig. lOD). Both tarsus I and II with pair of 
doubled pas. Medial pas on tarsus III setifonn; 
lateral pas spiniform, delicately pectinate. 
Medial pas on tarsus IV long and plumose, lateral 
pas short and bipectinate (Fig. lOB). 

Carpites on tarsi I and 11 8-9 long; carpites on 
tarsi III and IV 10 long. Accessory process on 
claws widened, about 4-5 wide, bearing 5-6 small 
tines. No tines on claw shaft. 

Female. Idiosoma 285-305 long. Outline of 
dorsal plates as in male though ornamentation 
lightly reticulate. Median portion of AD between 
ds-1 reticulate. Anterior margin of PD truncate; 
setae ds-4 at 0.2 1 -0.24. AE, PE and GP fused; this 
ventral shield separated from anal plate by 
narrow lateral wedges of striated integument. GO 
extending anteriad almost to level of insertion of 
leg IV. Area of genital plate with 5 pairs of pgs; 
two anterior pairs inserted distinctly anterior to 
GO (Fig. IOC). Genital sclerites with 2 pairs of 
sgs. Three pairs of tube-like genital acetabula. 
Genital spines claw-like. Tarsus III with 
spiniform lateral pas and 1-2 eupathid setiform 
medial pas (Fig. lOE). Lateral pas of tarsus IV 
similar to that of tarsus III; medial pas of tars us IV 
slightly smaller than lateral pas. 

Tritonymph. Idiosoma 248 long. Ventral plates 
separate; AE and PE each w ith a pair of adjunct 



ISO 



MEMOIRS OF THE QUEENSLAND MUSEUM 




F(G. 10. Rhombognathus levigatus Sp. nov.. A, larsus I. liucal. male (medial claw and setae omitte(0iB9tipOf 
t^sus IV, ventral, male (dorsal ^et^ptnUt^); C, idiosoma, ventral, female; D, tarsus 11, lateral, f emale (maiaL 
claw and setae omitted); E, tip.^tatsus 111^ ventral, female (dorsal setae omitted); F, posterior portiiprrc^ 
idiosoma, ventral, tritpnymphvasc^anal 5oJente^av=mial valve; gac^^g^ital^acetabula. Scal&i>af =ifi^uit^ 



:^;elae GP atid AP separate (Fig. I OF). GP with 2 
pairs of pgs and I pair of sgs. Telofemnra l-IV 
with 4/2. 4/2, 3/1. 2-3, ! dorsal/ventral setae. 
Setalion of the other segments same as in adults. 
Variatiahs. Varieties ofbfaaractEr!} in aOxtts: lengEh 
of idiosoma, female: 285-305 ( 5); length of idio- 
soma, male: 267-276 (3); number of adjunct setae 
on either side of AE: 1 (12), 2 (1); number of 
adjunct setae, PE: 1 (15), 2 (1); number of pgs on 
either side of GO, female: 5 (10); number of pgs 
plus basilar setae in cither halt male: 9+1 (2), 
10+1 (2), 11+1 (2); number of setae of leg 
segments 2 lo 5: 

leg II 

4/2(21. 
5/UI). 



segment 

a 
3 



leg! 

2(16) 
4/2(2), 
5/2(14) 



5(16) 
5(16) 



leg III 
-(16) 
3/i(i4j. 
4/Ul), 
5/UlJ 
3(16) 



leg IV 
2/1(4), 

mm 



5m 



Anomaly. In one of the females the left OC is 
lacking and replaced by striated mtegument. 

REMARKS. Rhombognaihun Uvtgstun is 

characterised by: smooth PD \\'ith single pair of 
setae; AE, PE and GP fused; area of AE and PE 
each with 1 pab of adjunct setae: female witli 5 
pmis Qip&\ male with 10-12 pairs of plumose 



telofemora MV with 5/2, 5/2, 3/1, V\ dorsal/ 
ventral setae. Slightly widened accessoiy process 

In the samples fi"ora the Great Barrier Reef, 
three species, Rhotvbo<^nathi4S levigafus, R. 
reiwuliff} sp. nov. and 7?. tericidits sp. nov., have 
claws with the shghtly widened accessor^' pro- 
cess bearing a few tines. In contrast to R. Jevigaius, 
the dorsal places oiR, r4ti(n4ifcr 9sx±S^ ter (cuius 
liave a distinct ret!icalat&eH|tutt^ ana long 
ds- 1 . Tlie three specif. di£E^HD the number 
of dorsal/ventral setae on the telofemora I to IV. 

Compared io Rhomhogiiathus species from other 
parts of the world, R. levlgatus is siinilar to R. 
notopsoides Bartsch, 1979, R, semir^ticufatus 
Bartsch, 1977, R. slnemfs BartsCh, IwO, andjft 

ventralLs Newell. I9S4. R nofopsoides is a 
brackish water species tYom eastern North 
America (Bartsch. 1 "^V^b). R sinensis is recorded 
from southern China and Japan (Bartsch, 1^*^0; 
Abe I996)> and R- sennrcticulatus and 
v^^^Uf&oni the £afit^ Pacific tBartjsQb, I97.7i: 
"NeweO, 1984). The dors»f plates bf S nof<h 
psoides are distinctly ornamented. In R. 
semireficidafifs, too, each of the dorsal plates 
bears .i d! -:n!-;i ■ i^iiucntation, and, in coiuraslto 
R. tevigatus^ each of the telofemofa 1 and Uliasi 
dorsal setae Imiohly 1 ^^mAts^la'R.iS^emk^ 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



181 



the arrangement of the setae on the telofemora I 
to rV is the same as in R. levigatus but the outhne 
and the ornamentation of the PD is different, and 
the number of tines on the accessory processes is 
somewhat larger than in the latter species. The 
telofemora I to IV of /?, ventralis bear 6, 6, 4, 4 
setae, the ds-1 are rather long, whereas in R. 
levigatus the ds-1 are short and the telofemora 
bear 7, 7, 4, 4 setae. 

Rhombognathus longipes sp. nov. 
(Figs 11-13) 

ETYMOLOGY. From (Latin) longus, long, and pes, foot, 
leg, for the long legs. 

MATERL\L. HOLOTYPE. 6 (MTQ), Great Barrier 
Reef, 18°25.93'S, 1 47^ LirE,Faradav Reef, coarse sand 
and mbble, lOm, 13 April 1998; J.C. Otto. PARAnTES. 
39, 2 c^, 1 proton\mph (MTQ), collection data as above. 
9, S (OM S50966), collection data as above. 2 9 (WAM 
99/ 1 445, 1 446), collection data as above. One 9 , S (ZMH 
A 1 00/99), collection data as above. 6 9 , 2 (5 , 1 tritonymph, 
2 deutonymphs (IB), collection data as above. OTHER 
MATERL^L. S (MTQ), Great Barrier Reef, I8"25.93'S, 
1 47°2 LITE, Faraday Reef, coarse sand and rubble, 2m, 1 3 
April 1998; J.C. Otto. 

DESCRIPTION. Male. Idiosoma slender, 317-328 
long; holotype 335 long, 185 wide. Dorsal plates 
with delicate reticulation. AD 11 2 long, 100 wide; 
anterior margin broadly rounded; posterior margin 
ovate. Line of internal scars at about 0.75 (Fig. 
11 A). OC 100 long, 55 wide; anterior cornea 
slightly larger than posterior one. Distance 
between gland pores 50; pore canaliculus almost 
halfuay between gland pores. PD 167 long, 97 
wide, 1 .7 times longer than wide. Pair of post- 
erior cones of PD almost extending to end of anal 
cone; each cone with gland pore. Setae ds-1 
10-15 long, positioned in posterior half of AD at 
0.64, i.e. somewhat anterior to transverse line of 
scars. Second pair of setae on OC almost at 0.43. 
PD with single pair of setae at 0. 1 6. Adanal setae 
distally on anal plate. Anal sclerites well developed. 

AE, PE, GP, and AP ftised (Fig. 1 1 B). Area of 
AE with pair of adjunct setae, that of PE with 
and 1 adjunct seta. GO 35 long, 25 wide; anterior 
margin level with insertion of leg IV. With 9 and 
10 pgs on either side of GO; pair of basilar setae 
incorporated in line of pgs (Fig. 1 IC). Spermato- 
positor 52 long, 46 wide, slightly extending 
beyond GO. 

Gnathosoma 90 long, 67 wide, 1.3 times longer 
than wide. Rostrum slender, apically pointed 
(Fig. 1 ID); almost as long as gnathosomal base. 
Narrow tectum truncate. Basal pair of maxillary 



setae in middle of rostrum; apical pair almost as 
long as basal pair. Slender palps appressed to 
rostrum. 

Legs slender; the four pairs similar in length 
and approximately 0.9 of length of idiosoma. 
Legs ill and IV at 0.53 and 0.69, respectively. 
Telofemora I-IV 2.7, 2.9, 2.7, 2.5 times longer 
than high. Legchaetotaxy: leg 1, 1, 2, 5, 5, 5, 3; leg 
II, 1 , 2, 5, 5, 5, 3; leg m, 1 , 1 , 3, 3, 5, 4; leg IV, 0, 1 , 
3, 3, 5, 3. Telofemora I-IV with 4/1,4/1, 3/0, 3/0 
setae. Two basidorsal setae on telofemora I and II 
short, spiniform (Figs HE, F). Bipectinate setae 
on tibiae I-IV numbering 2, 1, 1,2; these setae on 
tibiae I, II and III conspicuously long; on tibia IV 
ventrolateral bipectinate seta distinctly longer 
than ventromedial one. Tarsi slender. Basal setae 
of tarsi 111 and IV inserted in apical half of these 
seginents (Figs 12A, B). Two basal setae on 
tarsus III adjacent. Tarsus I slender, with elongate 
papilliform famulus and solenidion 14- long. 
Tarsi 1 and II with doubled pas; on tarsus III 
medial pas setiform, lateral pas short, spiniform 
(Fig. 1 2D); on tarsus IV medial pas long, plum- 
ose; lateral pas flattened, pectinate (Fig. 12E). 

Carpites on tarsi I and II 1 long, those on tarsi 
III and IV 11 - 1 2 long. Claws short; rounded apex 
with small accessory process. 

Female. Idiosoma 335-365 long. Dorsal aspect 
similar to that of male. AE, PE and GP fused. AE 
with 1, rarely 2 adjunct setae on either side; PE 
with 1, rarely 0, adjunct seta. GO 85 long; genital 
sclerites each with 2 sgs. With 5 pairs of pgs (Fig. 
12F). Two anterior pairs of pgs anterior to level of 
anterior edge of GO; anteriormost pair of setae 
slightly anterior to level of insertion of leg IV; its 
distance to edge of GO equalling 0.3 times length 
of GO. Posterior pair of pgs distinctly removed 
from the other setae. C^vipositor in rest reaching 
beyond GO (Fig. 12F). Extended ovipositor long; 
with basal pair of conical papillae and 5 pairs of 
apical genital spines (Fig. 12G). Genital spines 
8-9 long, each with 4 lateral tines. Gnathosoma 
with slender palps (Fig. 12H). P-4 with setae in 
basal whorl as illustrated (Fig. 121); tip with 2 
spurs and I setula. On tarsus IV medial pas 
shorter and less plumose than in male; lateral pas 
flattened and pectinate (Fig, 12J). 

Tritonymph. Idiosoma 340 long, 185 wide. 
Posterior margin of AD truncate (Fig. 13 A). OC 
short, only slightly extending beyond posterior 
gland pore; ds-2 within or just anterior to margin. 
PD much shorter than in adults. Ventral plates 
AE, PE. GP, and AP separated. Posterior portion 
of AE with projecting triangular or obtuse 



182 



MEMOIRS Of THE QUEENSLAND MUSEUM 




FIG. 1 1 . Rhomho^nathus longipes sp. nov., male; A, idiosoma, dorsal; B, idiosonui. ventral; C, posterior portion 
of idiosoma. ventral: D, gnathosoina. ventral: F:. leg I. medial; leg II, medial. Scale bar 50|jLm. 



portion, similar as figured in prolonymph (Fig, 
13E). AE and PE each witfi I pair of adjunct 

setae. GP w ith 2 pairs of pgs and I pair of sgs; 3 
iniiuile pairs of internal eenilal acetabula {Fiu. 
r^H). Teiofcniora 1-IV with 4/1, 4/1, 2/0, 2/0 
setae. Setation ot tlie other leg segments same as 
m adults. 

Deutt)nymphAdio$omR 236-263 kMig. Iti dorsal 

aspect similar to trilonymph. GP and AP fused 
(Fig. 13C); plate with 2 pairs of minute internal 
genital acetabula. 



Protonymph. Idiosoma 178 long, 105 wide. AD 
posteriorly trmcate; PD short, ovate (Fig. 13D). 

AE with 3 pairs of setae (Fig. 13E): PE with 1 
dorsal and 1 venlral seta. Genua 1-lV with 4, 4, 3, 
3 setae; telofemora l-III with 2/1, 2/1, 2/0 setste, 
femur IV with 2/0 setae. 

Variations, Varieti^ of characters in adults: 
length of idiosoma, 335-365 (10); length of 
idiosoma. o; 317-328 (8); number of adjunct 
setae on either side of AE: 1 (28), 2 (11); 3 (1); 
numb«' of dAysxi(A setae, PE: (5), 1 (35); 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



183 





FIG. 12. Rhombognathus longipes sp. nov.; A, leg HI. medial, male; B, leg IV, medial, male; C, tip of tarsus I, 
lateral, niaie (medial claw and setae omiilcd i; D, tip of tai'sus III, lateral, male; H, tip of tarsus IV, ventral, male 
(dorsal setae omitted): F, idiosoma, ventral, female; G, ovipositor, ventrolateral, female (spine in dotted line 
broken); H, gnathosoma, lateral, female; 1, P-2 ta P-4, lateral, female; J, tip of tarsus IV, ventral, female (dorsal 
setae omitted). Scale bar = 50|xm. 



number of pgs on either side, 9 : 5 (20); niunber 
of pgs on either side, d: 8 (8), 9 (7), 10 (1); 
number of setae of leg segments 2 to 5: 



segment 

2 
3 

4 

5 



leg I 

2(40) 
4/1(40} 

5(40) 
5(40) 



legll 

2(40) 

2/UI). 
4/1(36), 
5/1(3) 

m) 

5(40) 



leg III 

1(40) 
2/0(2), 
3/0(38) 

3(40) 
5(40) 



leg IV 
1(40) 

2/0(1), 

"1/0(30) 

5(40) 



184 



MliMOlRS OF THE QUJiENSLAND MUSliUM 




FIG. 1 3. Rhombo^nafhus Inu^/pes sp. no\ .; A. idiasoma, dorsal, Critonymph; B, posterior portion, of idiosoma, 
ventral, iritonyniph; C. posterior portion oi idio^oma, ventral, ifeutor^inph; D,idtosamatdOrsaliPralQnymph; 

L, idiosonia. veniral. prolon> mph. Scale bar ^ >OjjLm, 



separated Uncimcoiigeneiic Amtraltan specie^ on 

the basis pf the slender idiusoma with long, 
slender legs, 1^ I being almost a.s Iohl: as the 
idiosoma, J?, longipes is most sin.iij [o 
longisetus Bartsch, 1999, a species- toown trom 
New C^edonia. The most marlced di£fefence h 
tiMrlengthoflheds-l — in A longipes shatt^h^s 
•than 1/4 of the lengtfi of the AD, in R. lon^isetits 
as long as the AD. Apart from few exceptions, 
e.g. the ds-5 in the Mediterranean Copidognathm 
gwbus (Trouessart, 1889) and C. majusculiis 
(Tro|]«ss^^ the ieo|;UL of a seta is 

generally a ^stable character widiih a species. 
Fui'ther distinguishing characters are: The 
position ct the ds-I itn A*, fcngipes inserted at 
0.64, in R. longtscm:; ;ii < j ^7), the iengih of the 
ovipositor (in R. iongip^ distinctly eati^iulmg 
beyond the GO, in R/hf^ms Stiy sli^ti^ 
surpassing the G0>, the arrangement of the pgs 
around the female GO (in R longisetus almost 
equidistant whereas m R. Icmgipc^ the posterior 
pair of the setae is distanced from the preceding 
pair). The male GAof A longisetus has a post- 
genital papilla which iS lacking in R. longisetus. 

Rbombognatbus papuensis Baitsdi, ]9S9 

(Figs 14, 15) 

Riuwtbognuihus papuensis B^&ch, 1989a: 236, %s 5ii-f>5 
not HhombWMtht*^ pwutfnsis • ChsltiBriee, 1995. 282-284> 
figsi447 

MATERIAL. 9. 6 (NH Q), Great Bairier Reef Magnetic 
Island, Akaa, jq^ Uttota^ algae at (I5nv 16 March 



as above. 9. 6 (MTQ), Great Banier Reef 18°4l.29'Sj 
14T^5.83'E, Loadstone Reel; Halimeda at 3-6m, ) 1 April 
l<^98; coll. J.C. OOp. 9, i (QM S5096^,.CQ||f?^daia. 
i\s be&SQ. 49, 1 tritonymph(IB),col]e(iiQ0d^a^ 

1 <3i«at Biimer Reef, 19^0.1 rs, 

149TO.85^, EBzmdxIteeS UtAkmda sp. {Qiloropliyta) 
4 10ni,25 December 1997; coll. LC. Obd. 

DESCRIPTION (based on specimens from ^laUow 
water habitats from Magnetic Island). Idiosomal 

length of female 310, of male 277-285. Dorsal 
plates with foveate sculpturing. Posterior AD 
broadly rounded; foveate areolae d&tinCA'$)Lai:^a 
antmpr. to short L, OC larg^, ^tb % fxas^ 
aitd 2 ^unt set&e: 2 gland pOres attd oiiie 
canaliculus in lateral margin. PI^ with pair of 
wide costae; foveate areolae lateral and medial to 
costae (Fig. 14A). PD with single pair of setae, in 
females inserted at 0.24, in males at 0.3 1 . Ventral 
plates AE. PE and OA fiaed. On either side of AE 
and PE (0-)l adjunct setae Female with pair of 
lateral wedges between GPand AP;5 pan^ofpgs 
inseded almost equidistant ( Fig. 1 5F). Male with 
9- 1 3 plimiose pgs arranged in a line on either side 
of GO, and pair of basilar setae adjacent to 
pQsteapr part o£GO jpFig^ 14B). Sp^nnatapoator 
75 (ong; w wide, cJrtenditTg firlaeydnd GO (Fig. 
14C) 

GnathosomLi short; lcngth:\vidth ratio 1 .1 (Fig, 
15A). Rostrum conical, short, hardly more tim 
1/3 of length of gnalhosoma. Basal pair of maJc- 
illary setae longer than apical pair. Palps short. 

Legs I and II slightly shorter than legs III and 
IV. Telofemora I lo IV approximately 2,4 limes 
longer than high. T^lotftmora I and II longer than 



RHCftffiQGNATHllMAE; FRQM THE GREAT BARRIER REEF 



I8S 




PIG. \4,^hambognatfmspcipuertsiA Barl3ch,male: A, idiosoma, dorsal; B, idiosomd, venlraJ; C, posterior portion 



telofemora 111 and IV (Pigs 15B-E). Tibiae I-IV 
altnost equal in length. Tarsi I and II about as long 
as these legs' tibiae; tarsi III and IV distinctly 
longer than iibiae. Leg chaelotaxy (rare variants 
in parentheses): leg 1.1,2. (4- )6. (4- )5, 5, 3; leg II, 
U2.(5-)6, 5,5,3;leglll, 1,2,3.3,5,4; legIV,0, 
2, 3(.-4J. 3, 5, 3. Telofemora I-IV with Q/U 3/2) 
4/2, (3/2) A/2, m, 3/0 dGrsalArtniraf sctee. 
Ventral seta on genu I slender. Tibiae l-IV with 2, 
IJ , 2 bipectinate setae. Two basal setae on tarsus 
III inserted adjacent- Two disial fossar\ ^eiae on 
all larsi equal m size and barbate. Tarsus I with 
papillitorm famuluj&^dsl^Qdejf; i$eti&rm siblen- 

idion (Fig. 15G). 

Claws widened; truncate edge of this portion 
17 wide and provided with i s 2U uncs Apical 
endofclarw Separated trpm tnineaie wtdejied 
po!rth«i{Ffe. fSH). 

Varialions. Varieties of characters in aduhs; length 
of idiosoma, i : 310 (2), length i)f idiosoma, d: 
277-285 (2); number of adjunct setae on either 
side ofAE: 0(1), 1 (7k number of adjwot setae, 
PE: (1 ), I (6); number ofpgs on eithei'sidd Hf 
CO. ? : 5 (4); number of pgs plus basilar setae in 
cuhei halt: i\ 9+1 (1), 12+1 (2), 13H-1 (1); 
number of setae of leg segments 2 to 5: 



depth, respcctivel) , differ slightly from the above 
outlined characters. Females are 204-275 long 
and males 223-241 long. The dorsal plafefs 'are 
more distinctly reticulated. The PD is slightly 
more slender; its cosiae are distinctly separated 
from the reticulate remainder The number of pgs 
on the n^e GP is slightly smallef. 13ie widened 
trtmCateedge of ^tistJaws we 12 iWde. 

Variation iti the number of setae in specimens 
from the Elizabeth Reef and Loadstone Reef is as 
follows: number of adjunct setae on either side of 
AE: 1 (17J;nimiberofadjunctsptaeQnF£: 1 (17)^ 
d{0; tittrtiber of pgs oneith^aidcOTGO, female: 
5 (6); number ofpgs on either side plus basilar 
setae, male: 9-hi (5), 10+1 (I), number of setae of 
l%5egnijBi05 24)DL5: 

>niimt tegJ iegil legxtl leg IV 

t mm ^w) ais) i(2),2ae} 



3 



5(17) 
H 17) 



4/2(18) 

5(18) 



3/0(l2J 
3(17) 
5(17) 



3/0(17) 
3(18) 



segment 




icg U 


leg 111 


leg [V 


2 


W) 


2(8) 


2(8) 


2(8) 


3 


Ivll 1 1, 


3/2{3), 
4/2(4) 


3/0(S) 


3/0(8) 


4 


4tt),5C7) 


5(8) 




3|8I 


5 


5f8) 




5(8) 


518) 



REMARKS. The specimens from the Elizabeth 
Iteef 3nd Li>adst6tie Re^ )0m and3^ 



The tritonymiAs irom Elizaibetli Reef ^md 
Loadstone Reef ^6 chafactetfsed by! ldic>S5iia& 

205-229 long. Ventral plates separated. GP 
separate from AR AE with 1 pair of adjunct setae; 
PF with 0- 1 adjunct setae. GP with 2 pairs of pgs, 
1 pair of sgs. Leg chaetotaxy from trochanter to 
tarsus (rare variants in parentheses): leg I, 1,2, 
(4-)5.5.5,3;legII 1- 2, (4-)5. 5, 5, 3; legin, 1,2, 
2, 3, 5, 4; leg FV, 0, 2. 2, 3, 5, 3. Telofemora I-IV 
with (2/2) 3/2, (3/1) 3/2, 2/0, 2/0 dorsal/ventral 
setae. Two basal setae pa tarsus ill actjacent. Edge 



186 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 15. Rhombognathus papuensis Bartsch; A, gnathosoma, ventral, male; B, leg 1, medial, male; C, leg II, 
medial, male; D, leg III, medial, male; E, leg IV, ventromedial, male; F, idiosoma, ventral, female; G, tarsus I, 
lateral, female (medial claw and setae omitted); H, tip of tarsus III, medial, female. Scale bar = 50|jLm. 



The specimens from the Great Barrier Reef 
differ slightly from those from Papua New 
Guinea (Bartsch, 1989a). In the specimens from 
Papua, the setae on the PD insert further 
posterior, and the wedges between GP and AP 
seem to be larger than in adults from the Great 
Barrier Reef Unless more material will prove the 
opposite, the individuals from Magnetic Island, 
Elizabeth Reef and Loadstone Reef are con- 
sidered as conspecific with R. papuensis. 



The individuals recorded from the Indian Ocean 
(Chatterjee, 1995) are not conspecific with 
Rhombognathus papuensis. According to the 
description of that species, the OC are much 
wider than in R. papuensis and the PD bears 2 
pairs of setae (Chatterjee, 1995: Fig. 1). 

DISTRIBUTION- New Guinea, 10°S, 148°E, 
shallow water (Bartsch, 1 989a) and Great Barrier 
Reef, from shallow water to 10m depth. 



RJIOMBOGNATIIINAE FROM THE GREAT BARRIER REEF 



187 



Rhombognathus reticulifer sp. nov. 
(Figs 16, 17) 

ETYMOLOGY. For the dorsal plates which bear iferre, 
Latin) a reticulum (Latin). 

MATERIAL. HOLOT\'PR. S (MTQ), Great Barrier 
Reef, I9°20.12'S, 149°02.85'E, Elizabeth Reef. large 
chunks of coral rubble ai 10m, 24 December 1997; coll. 
J.C. Otto. PARAT\TES. 29 (MTQ), collection data as 
above. 9, S (QM S50969), collection data as above. 
\6 (ZMIl AlOl/99), collection data as above. 6 (IB), 
collection dam as above. OTHER MATERIAL. 79,25,2 
tritonymphs (IB). Great Barrier Reef, 19°20.12'S. 
149°02.85'E, Elizabetli Reef, Halimeda (ChlorophMa) at 
1 5m, 24 December 1997; coll. J.C. Otto. 

DESCRIPTION. Mate. Idiosoma slender, 186- 
205 long; holotype 203 long, 132 wide. Dorsal 
plates coarsely reticulated (Fig. 1 6A); meshes 8- 1 
long and faintly subdivided. Plates with delicate 
pores. AD 62 long, 65 wide. Anterior margin 
arched, posterior margin broadly rounded. Trans- 
verse series of muscle scars level with 0.80. OC 
54 long, 30 wide. Each plate with 2 small comeae, 
2 gland pores in lateral margin and pore canal- 
iculus half\vay between gland pores. PD 105 
long, 62 wide; not as wide as AD. Plate evenly 
reticulated; meshes 8-11 long. Posterolateral 
portions of PD hardly projecting beyond median 
portion of plate. Pair of gland pores in postero- 
lateral margin of PD. Anal scleriles extending 
bevond iuial valves. Setae ds-1 40 Ions, inserted on 
AD. Setae ds-2 and ds-3 on OC; ds-3 at 0.51. PD 
with single pair of setae, inserted at 0.25 and 0.29. 

Ventral plates AE, PE, GP and AP fused to a 
ventral shield (Fig. 16B). Shield delicately 
punctate. Area of AE with 1-2 adjunct setae; PE 
w ith 1 pair of adjunct setae. GO 28 long, 1 6 wide. 
GO extending to the level of insertion of leg IV. 
Perigenital setae arranged trapezoidally; holo- 
type with 7 pairs of setae in a line and 1 pair of 
basilar setae near posterior edge of GO. Pgs 
plumose. Spermatopositor 37 long, 40 wide; 
extending beyond anterior pair of pgs (Fig. 1 6C). 

Gnathosoma short; 57 long, 46 wide; 1 .2 times 
longer than wide (Fig. 16D). Rostrum short, 22 
long, triangular. 

Legs 1 and IV almost equal in length, about 0.7 
of length of idiosoma. Insertion of legs III and IV 
level with 0.56 and 0.71, respectively. Length: 
height ratio of telofemora I-IV 1.7, 1.7, 1.5,^1.6 
(Figs 16E-H). Telofemora and tibiae of each leg 
almost equal in length. Tarsi III and IV only 
slightly longer than these legs' tibiae. Leg 
chaetotaxv, from trochanter to tarsus: leg 1, 1,2, 
6, 5, 5, 3; leg II, 1, 2, 6, 5, 5, 3; leg III, 1,1,3,3,5, 
4; leg IV, d, 1,3, 3, 5, 3. Lateral seta on each 



basifemur II and III long, about twice height of 
these segments. Telofemora I-IV with 4/2, 4/2, 
3/0, 3/0 setae. Dorsolateral seta on telofemur III 
hardly longer than the 2 dorsomedial setae. 
Ventral seta on genu 1 delicately pectinate. Tibiae 
I-fV with 2, 1, 1,2 bipectinate ventral setae. 
Ventrolateral seta on both tibia 1 1 and III almost as 
long as ventromedial seta. Two basal setae on 
tarsus III inserted close together. On each of the 
tarsi 2 distalmost dorsal setae slightly plumose. 
Tarsus I with short papilliform famulus, 1 long; 
solenidion 7 long (Fig. 17.^); ambulacrum 
flanked by pair of doubled pas (Fig. 17B). Apart 
from absence of famulus, tarsus II similar to 
tarsus I; solenidion 8 long (Fig. 1 7C). Medial pas 
on tarsus III setiform, lateral pas short, pectinate 
(Fig. 17D); pas on tarsus IV (Fig. I7E) more 
plumose than on tarsus IIL 

Carpites on tarsi 1 and II 5-6 long; carpites on 
tarsi 111 and IV 6-7 long. Accessory processes of 
claws widened; 4 wide, with 7-8 small tines (Fiss 
17D,E). 

Female. Idiosoma 217-241 long. Areas with 
striated integument between plates larger than in 
males. Feinalc PD somewhat shorter, ds-4 at 
0.20-0.23 relative to length of PD. Ventral shield 
including AE, PE and GP; AP separated from 
ventral shield by wedges of striated integument 
(Fig. 17F). Anterior margin of CiO not reaching 
the level of insertion of leg IV. Genital acetabuia 
small, often obscured. Five pairs of pgs arranged 
in a wide ring around GO; 2 anterior pairs of pgs 
anterior to the level of insertion of leg IV. Genital 
spines claw-like, with median tooth and 4-5 
smaller teeth along lateral margin. Genital 
scleriles with 2 pairs of sgs. Palps extending 
slightly beyond rostrum. Chclicera 70 long. 16 
wide; dorsal margin of its claw dentate. Pas on 
tarsus III similar to male tarsus; pas on tarsus IV 
(Fig. 1 7H) less plumose than on male tarsus IV. 

Tritonymph. Idiosoma 151-1^5 long. OC more 
narrow and PD shorter than in adults (Fig. 171). 
Reticulate pattern same as in adults. Setae ds-1 
long; setae ds-4 near anterior margin of PD. 
Ventral plates AE, PE, GP, and AP separated (Fig. 
17J). Posteromedian margin of AE convex. AE 
and PE each with 1 pair of adjunct setae. Genital 
plate small; with 2 pairs of pgs and 1 pair of sgs. 
Gnathosoma short; 1.1 times longer than wide. 
Legs I and U with 1, 2, 4, 5, 5, 3 setae, from 
trochanter to tarsus; leg III with 1, 1, 2, 3, 5, 4 
setae, and lee IV with 0, 1,2, 3, 5, 3 setae. 
Telofemora !-Tv with 3/1 , 3/1 , 2/0, and 2/0 setae. 

Variations. Varieties of characters in adults: 
length of idiosoma, 9: 217-241 (II); length of 
idiosoma, 6: 186-205 (6); number of adjunct 



188 



MEMOIRS OF THE QUEENSLAND MUSEUM 




¥lGA^.Rhpmb9^athusreti^liferSp. txm.f male; A, idiosoma, dorsal; B, idiosoma, ventral; C^posterior portion 
of idiosoma, ventral; D« giiatbosoma, v^tral; E, 1^ I, media]; basi&iitur io tatsos II, icffidia}; G, leg til, 
medial; H, leg IV, medial. Scale bar= 50(un. 



setaeoneithersideof AE: I (29), 2 (5); number of 
adjunct setaeonPE: 0(3), 1 (30), 2(1); niimberof 
pgs on either side of GO, 9 : 5 (20), 6 (2);numt)er 

of pgs plus basilar setae in either half, <?: 7+1 
(1 1), 8+1 (1); number of setae of leg segments 2 
to 5: 



segment leg I 

2 2(34) 

3 3/2(4), 
4/1(1). 
4/2(29j 

4 S(34> 

5 m) 



legll 
2(34) 
3/2(4), 
4/2(30) 

5(34) 
5(34) 



legm 
1(34) 
2/0(7). 
2/1(1). 
3/0(26)^ 
3(34) 
5(34) 



leg IV 
1(34) 

2/0(9), 
2/1(1). 
3/0(24) 
3(34) 
5(34) 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



189 




FIG. 1 7. Rhombognathus reticulifer sp. nov., A, tarsus I, lateral, male (medial claw and setae omitted); B, tip of 
tarsus I, ventromedial, male (dorsolateral fossary seta omitted); C, tarsus II, lateral, male (medial claw and setae 
omitted); D, tip of tarsus III, ventral, male (dorsal setae omitted); E, tip of tarsus IV, ventral, male (dorsomedial 
fossary- seta omitted); F, idiosoma, ventral, female; G, gnathosoma, lateral, female; H, tip of tarsus IV, ventral, 
female (dorsal setae omitted); I, idiosoma, dorsal, tritonymph; J, idiosoma, ventral, tritonymph. Scale bar = 50|xm. 



REMARKS. Rhombognathus reticulifer is 
characterised by the combination: dorsal plates 
distinctly reticulated; ds-1 nnuch longer than the 
succeeding setae; PD with single pair of setae; in 
males all ventral plates fused; females with 
wedges of striated integument between ventral 
shield and AP; area representing AE and PE 
generally with 1 adjunct seta on either side; males 
with 8 pairs of pgs, females with 5 pairs of pgs; 
gnathosoma short, 1.2 times longer than wide; 



telofemora I-IV with 4/2, 4/2, 3/0, 3/0 setae; 
accessory process on claws slightly widened, 
with 7-8 tines. 

Amongst the rhombognathines from the Great 
Barrier Reef, the species R. levigatus and R. 
tericulus sp. nov. (description below) are most 
similar to R. reticulifer. The smooth PD of R. 
levigatus is wider than in R. reticulifer, and the 
telofemora 1 to IV of R. levigatus bear 5/2, 5/2, 
3/1, and 3/1 setae. Discriminating characters 



190 



MEMOIRS OF THE QUEENSLAND MUSEUM 



between R. reticidifer and R. tericulns are 
outlined after the description of the latter species. 

R. ventj'alis Newell, 1984 and/?, lateralis NewelL 
1 984, both knowTi from the South American Pacific 
coast (Newell, 1984), resemble R. reticulifer in 
general aspect. These two species have, in 
contrast to R. reticidifer, 4 setae on telofemur IV. 

Rhombognathus scutulatus Bartsch, 1983 

Rhombognathus saitulatus Bartsch. 1 983: 4 1 3-4 1 5, ligs 46-57. 
Rhomhognathus scutulatus Bartsch, 1993: 20, 21, I1g. !A-C; 
Chalterjec, 1995: 284, figs 15-19. 

MATERIAL. 6 (MTQ), Great Barrier Reef, Cape 
Ferguson, AIMS beach, algae at low tide mark, 2 March 
1997; coll. J.C. Olto. 9, S (QM S50970), collection data 
as before. S (IB); collection data as before. 9, 6 (MTQ), 
Great Barrier Reef, Magnetic Island, Alma Bay, rocl^ 
littoral, algae at 0.5m, 1 6 March 1 998; coll. J.C. Otto. 2 9 , 
6 (IB); collection data as before. 

DIAGNOSIS. Female idiosoma 300-335, male 
285-310 long. Dorsal plates AD, OC and PD 
fused. Dorsal shield with foveate ornamentation. 
Dorsal idiosomatic setae subequal in size. Area 
representing PD with single pair of setae. Ventral 
plates AE, PE and CP llised in females and males; 
GP partly ilised with AP. Areas of AE and PE 
each with 1 pair of adjunct setae. Female with 5 
pairs of pgs and 2 pairs of sgs. Males with 9-13 
pairs of trapezoidally arranged plumose pgs, I 
pair of basilar setae and 2 pairs of sgs. 
Gnathosoina 1.16 times longer than wide. None 
of the setae on genua I-I V pectinate. Telofemora I 
to IV with 4/2,4/2, 3/0, 3/0 setae. Tarsi I-IV with 
3, 3, 4, 3 dorsal setae. Apical fossary setae with 
their distal portion flaUened, pilose. Accessory 
process of claws widened; claw with 22-24 tines. 
Variations. Varieties of characters in adults: 
length of idiosoma, 9: 300-335 (4); length of 
idiosoma, S: 285-310 (5); number of adjunct 
setae on either side of AE: ( I ), 1 ( 1 7); number of 
adjunct setae on PE: 1 ( 1 7), 2 ( 1 ); number of pgs 
on either side of GO, 9 : 5 (8); number of pgs on 
either side plus basilar setae, i\ 9+1 (2), 10+1 
(3), ll+I (2), 12+1(1), 13+1 (2); number of setae 
of leg segments 2 to 5: 



segment 


Icgt 


legll 


leg III 


leg IV 




2(18) 


:<18) 


2(18) 


2(18) 


3 


4/:(i8) 


4/2(17), 


2/0( I ). 


2/0(1), 




4/3(1) 


3/0(17) 


3/0(17) 


4 


5(18) 


5(18) 


3(17), 4(1) 


3(18) 


5 


5(18) 


5(18) 


5(18) 


5(18) 



REMARKS. In the samples from the Great 
Barrier Reef area, Rhombognathus scutulatus is 
the only species with a dorsal shield. 



DISTRIBUTION. Philippines; eastern and 
western coast of India; Western Australia 
(Bartsch, 1983, 1993; Chatterjee, 1995). Found 
in a variety of tidal and shallow subtidal algal 
substrata. 

Rhombognathus seminotatus sp. nov. 
(Figs 18, 19) 

ETYMOLOGY. For the sculpturing of the PD with half 
(semi, Latin) of the plate being ornamented (notatus. Latin). 

MATERIAL. HOLOTYPE. 6 (MTQ), Great Barrier 
Reef, Cape Ferguson, AIMS beach, algae at low tide mark, 
2 March 1 997; coll. J.C. Otto. OTHER MATERIAL. 9 , 6 
(MTQ), Great Barrier Reef, Magnetic Island, Alma Bay, 
rocky littoral, algae at 0.5m, 1 6 March 1 998; coll. .I.C. Olto. 
9, 6 (QM S5097I). collection data as above. 9 (ZMH 
A 102/99), collection data as above. 29, 2d (IB), col- 
lection data as above. 

DESCRIPTION. Male. Idiosoma slender 278- 
313 long; holotype 278 long, 161 wide. AD 100 
long, 82 wide. Anterior and posterior margin 
broadly rounded. Median portion with reticulate 
ornamentation; anterior, lateral and posterior 
portions smooth (Fig. ISA); lateral areas pierced 
by minute pores. Transverse line of muscle scars 
at 0.78. OC 75 long, 48 wide; with 2 corneae; 
gland pores in lateral margin. PD 130 long, 85 
wide. Posterolateral cones of PD hardly extend- 
ing beyond posteromedian margin of the plate; 
each cone with a gland pore. Median and 
posterolateral portions of PD reticulate; costae 
slightly raised, smooth apart from minute pores. 
Anterior portion of PD in holotype almost 
smooth; in other specimens that portion covered 
by delicate epicuticular reticulum (Fig. 18C). 
Anal valves not extending beyond anal sclerites. 
Setae ds-1 30 long, inserted on AD at 0.50. 
Succeeding setae 1 5 long; posterior seta on OC in 
posterior half of the plate. PD with single pair of 
setae at 0.28. Adanal setae on anal valves. 

Ventral plates AE, PE, GP, and AP fused to a 
ventral shield (Fig. 18B); shield 218 long. Areas 
representing AE and PE each with a pair of 
adjunct setae. GO extending anteriad far beyond 
the level of insertion of leg IV. GO 45 long, 21 
wide. Distance to end of anal cone equalling 
length of GO. Perigenital setae arranged 
trapezoidally, 12 and 14 plumose setae in line and 
1 pair of basilar setae at the level of 0.8 relative to 
length of GO (Fig. 1 8D). Genital sclerites with 2 
pairs of sgs. Spermatopositor 62 long, 62 wide; 
extending beyond anterior pair of pgs. 

Gnathosoma short, 71 long, 57 wide; length: 
width ratio 1 .24 (Fig. 1 8E). Tectum slightly arched 



RKOMBOGNATHINAH f'KOM THE GRiiAT BARRIER REEF 



191 




FIG. 18. RhombogfiathiLs seminoratus sp. nov.. male; A^, idiosoma, dorsaJ; B, idiosoma, vejiiral; C, anterior 
portion of PD; D, posterior portion of idiv)soma. ventral; E, gnalhosoma, ventral; F, gnalhosoma, lateral; G, 

ledum and P-I to P-3. dorsal. Scale bar -- 5i)\im. 



(Fig. J SG). Rostrum 25 long; apex poinied. Palps 
short, oppressed to the rostrum. P-4 directed 
Vfirttrad (Fig. 1 8F), generally obscured in dorsal 
aspect (Fig. 18G). 

Legs approximately 0.8 limes of lenulli of 
idiosoma. Insertion of leg I at 0.12. that of leg IV 
at 0.72 relative to length of idiosoma. Telofemora 
1-IV 2.5, 2.4, 2.3, and 2.3 limes longer than high. 
Tdpfemgraiand il markedly longer than tibiae 1 
andftrespecfivety (Fig. t9A, B), telofemora 111 
and IV somewhat longer than these legs, tibiae 
(h'igs l^JC. D). Tarsi 1 and 11 approvimaiely as 
long as tibiae land II; larM ill .duI IV longer than 
fibiae. Leg chaetotaxy: leg i. 1, 2-3, 5-(), 5, 5, 3; 
legU, 1, 2, 6, 4-5, 5, 3; leg IIT, 1 , 2, 3-4, 2-3, 5, 4; 
leglV, 0, 3, 3, X 3. None of ventrolateral setae 
on gemia bipectmale. Tibiae I-IV with 2, I, 1,2 
bipectmate setde. Qj^ tibia IV v^dtroinedial seta 



smaller than ventrolateral one. Paired dorsal 
setae of tarsi slightly plumose.. Tkrsus III with 4 
dorsal setae; the 2 ba^ gne^ insecte^ ^i^j^geot- 
Solenidia on tar^i I and ?1 9 and 11 Idtig, 

respectivelv. Famulus on tarsus 1 papilliform, 2 
long(Fig. IQF ). Tarsi 1( Fig. 19F) and 11 each with 
pair of doubled pas. Medial pas on tarsus 111 
setifonrL, lateral pas spinifortn, btpectiiiate. Medial 
pas on tarsus Ivtong-, phlmcse (Fig, 19H). 

Tarsi 1 and II each with 9-10 long carpite; tarsi 
III and IV with 12 long carpj^. Claws distally 
widened; with 12^13 tw^-End-of^law clearly set 
off from line wiflt ftnes. 

Female. Idiosoma 3 1 0-322.^triate(l integument 
between dorsal plates wider thsttl initiales. AE, 
PE and CiP tlised to a ventral shield. Nan\»w 
striated integument separates anal plate from 
vetilfal s(hield. GO 67 long, 45 wide; extending to 



192 



MEMOIRS OF THE QUEENSLAND MUSEUM 



the level of insertion of leg IV. Seven to nine pairs 
of pgs arranged in a wide ring aroimd GO (Fig. 
191); anterior pair of pgs level with insertion of 
leg IV. Genital scleriles with 2 pairs of sgs. 

Variations. Several individuals from Magnetic 
Island have the dorsal plates covered by an 
epicuticular reticulum, the reticulum formed by 
delicate droplets (Fig. 18C). 

Varieties of characters in adults: length of idio- 
soma, ?: 310-322 (4); length of idiosoma, i: 
278-313 (5); number of adjunct setae on either 
sideofAE: 1 (20); number ofadjunct setae on PE: 
0(3), 1 (17); number of pgs on either side of GO, 
female: 7 (3), 8 (4), 9 (3); number of pgs plus 
basilar setae in either half, male: 10+1 (1), 11+1 
(5), 12+1 (2), 13+1 (1), 14+1 (1); number of setae 
of leg segments 2 to 5: 

segment leg I leg II leg Hi leg IV 

2 2(17), 3(1) 2(18) 2(18) 2(18) 

3 2/1(1), 4/2(18) 



2/1(1), 
4/1(1), 
4/2(15) 

5(18) 
5(18) 



4(1), 5(17) 
5(18) 



2/0(3). 2/0(1), 

3/0(14), 3/0(i7) 
3/1(1) 

2(1). 3(17) 3(17). 4(i; 

5(18) 5(18) 



REMARKS. Rhombognathus seminotatus is 
characterised by the distally widened claws 
bearing 12-13 tines. Other characters are: AE and 
PE each with pair ofadjunct setae; females with 
7-9 pairs of pgs, males with 11-15 pgs; telo- 
femora I-IV with 4/2, 4/2, 3/0, 3/0 setae. In the 
majority of species the 2 basal setae on tarsus III 
insert immediately adjacent. 

The majority of congeners of the Great Barrier 
Reef area have claws which are smooth or have 
an accessory process with 0-8 lines, other species 
have distally widened claws with more than 15 
tines {R. papuensis, R. scutidatus), whereas R. 
seminotatus has 12-13 lines. Widened claws are 
present also in the eastern Pacific species R. atuv 
Abe, 1990, R. guamensis Barlsch, 1989, and R 
sinensoideus Bartsch, 1992, recorded from 
Hokkaido, Guam, New Guinea, and southern 
China, respeclively (Abe, 1990; Bartsch, 1989a, 
1992). In contrast to R. seminotatus, the PD of 
both R. atuy and R. sinensoideus has 2 pairs of 
setae. The claws of R. guamensis have unusually 
wide lines, evidently wider than in R, semi- 
notatus. R. insidaris Bartsch, 1989, a species 
from islands of the Hawaii Archipelago (Bartsch, 
1989a) and R. lateralis Newell, 1984, recorded 
from South America (Newell, 1984), also have 
claws similar to those of R. seminotatus. The 
three species can be distinguished on the basis of 
the setation of the telofemora I to IV; in R. 
insular is there are 5/2, 5/2, 3/1,3/1 dorsal/ventral 



setae, in R. seminotatus 4/2, 4/2, 3/0, 3/0 setae, 
and, according to Newell (1984), the setal 
formula of the telofemora 1-lV in R. lateralis is 6, 
6, 3-4, 4. 

Females of i?. seminotatus can be rapidly dis- 
criminated from congeners on the basis of the 
combination of: GAwith 7-9 pairs of pgs; AE, PE 
and GP fiised to a ventral shield; and claws with 
approximately 12 tines. The other rhombognathine 
species from the Great Barrier Reef area have 5 
pairs of perigenital setae. A high number of peri- 
genital setae, similar to that in R. seminotatus, is 
present in the Subantarctic species R. auster 
Bartsch, 1989b and/?, c/anvm/ Newell, 1984, the 
eastern Pacific R. eUipticus Bartsch, 1977, and 
the northeastern Atlantic R. procerus Bartsch, 
1 975b, but none of these species have claws with 
a widened accessory process bearing more than 
10 tines. 

Rhombognathus tericulus sp. nov. 
(Figs 20,21) 

ETSTVIOLOGY. Tericulus, an anagram of reticulus which 
refers to the reliculate ornamentation of the dorsal plates. 

MATERIAL. HOLOTYPE. 6 (MTQ), Great Barrier 
Reef, 18°25.25' S, 146°40.65'E, Bramble Reef, chunks of 
coral rubble, 3-6m, 10 April 1998; coll. J.C. Otto. 
PARATYPES. 2, 6 (QM S50972), collection data as 
above. 26 (MTQ), collection data as above. OTHER 
MATERIAL. 9 (MTQ), Great Barrier Reef, 19°20.12'S, 
149°02.85'E, Elizabeth Reef, coarse sand and rubble, 3m, 
25 December 1997; coll. J.C. Otto. 6 (QM S50973), same 
collection data as before. 6 (WAM 99/1447), same 
collection data as before. S (IB), same collection data 
as betbre. 29, c?, I tritonvmph (IB), Great Barrier Reet^ 
18°41.9rS, 147m.49'E,^ Loadstone Reef, coarse sand 
and nibble at 2m, 12 April 1998; coll. J.C. Otto. 9 (IB), 
Cireat Barrier Reef I8°42.05'S, 147°05.98'E, Loadstone 
Reef, coarse sand and rubble, 8m, 12 April 1998; coll. J.C. 
Otto. 29 (MTQ), Great BanierReef, 14°36'S, 145"38'E, 
Yonge Reef, coarse sand and rubble, 9m, 1 October 1 998; 
coll. J.C. Otto. 9, 6 (ZMH A103/99), collection data as 
before. 2 9 , (IB), collection data as before. 

DESCRIPTION. Male. Idiosoma 205-223 long; 
holotype 220 long, 143 wide. Dorsal plates 
conspicuously reticulated (Fig. 20A). AD 75 
long, 75 wide; reticulation of anterior portion of 
plate indistinct; posterior line with muscle scars 
inserted at 0.72. OC 68 long, 42 wide; distinctly 
projecting beyond posterior gland pore. PD 127 
long, 82 wide; prominent reticulation uniformly 
covering the plate. Posterior margin of PD not 
extending beyond anal cone. Setae ds-I very 
long, appro-ximately 40; inserted in anterior half 
of AD and close to the level of gland pores. 



RHOMBOGN ATHINAE FROM THE GREAT BARRIER REEF 



193 




riG. 19. Rhomhognaihus scmimnants sp. nov.; A. leg L ventromedial, male; B. leg II. vcntroniediaL male; C, leg 
III, mediai. male; D. leg IV, medial, male; E, tip of tarsus I, ventral (dorsal setae omitted); F, tarsus 1. lateral, 
male (medial claw and setae omitted); G, tarsiis 11, lateral, male (medial claw and setae omitted); H. tip oftai'sus 
IV^ miedial, male (lateral setae and ^law dashed); 1, idiosoma, ventral, female. Scale bar - SOp.ra. 



Following setae 10-12 long. Setaed^^ en ?D at 
0.37. Adatidl setae on 

Ventral plates AE,PEftnd GAllised (Fig. 20B). 
Area of AE and PE eacli with one pair of adjunct 
setae. Ntoginal setae long. GO 30 long, 1 9 wide; 
distaac9to of ao^ CQne saore than length of 
GO. On either side of GO Mne with 8 plumose pgs 
and 1 basilar seta; the latter inserted anterior to 
posterior end of GO (Fig. 20E). Spermatopositor 
32 long, 39 wiije. 



Gnatfidsoma short, 65 long, 52 Wide. Rostrum 

27 long. Tectum truncate (Fig. 2()C). Chelicera 70 
long; its claw wide, cutting edge serrate (Fig. 
2DD). Palps^eit. 

Four pairs of legs subequal in length and 
^proximately 0.7 of length of idiosoma.Leg3 lU 
and IV infeeited at 0.56 and 0.71, respectrvely. 

Teiofemora I-T\' 1 .3-1 .4 times longer than high. 
Teiofemora i and 11 shghtly longer than tibiae of 
these leg^; teiofemora 1U and IV sKghtly shorts^ 



194 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2f^, JUiombognaihus tericulus sp. nov., male; A, idiosoma, dorsal; B, Miosoma, ventral; C, gnathosoma, 
lateral; D, tip of cheiicera; E, posterior portion of idiosoma, ventral; F, leg I, ventromcdiaT; G, leg II, 
ventromedial; H, leg HI, ventral; 1, leg IV, -ventral. Scale bar = SO^un. 



than tibiae. Leg chaetotaxy, from trochanter to 20F-I). Tibiae I-IV with 2, 1, 1.2 bipectitmte 

tibia: leg U, 2, 5, 5. 5, 3; leg IIJ, 2, 6, 5- 5, 3: leg ventral setae. Distal pair of fossary setae 

111, U 1, 4, 3, 5, 4; leg IV, U, 1, 4, 3, 5, 3. delicately plumose. As in female, soienidion on 

Teloferaoral-rVwith4/l,4/2,3/U3/lsetae(Fig. tarsus I setifonn, 7 long; feimilus 1 long (Fig, 



RI luMBl KrNA n UN AK FROM THE G^¥JiT BARRIIiR RKHF 



- 1 1: ). Tarsi I and 11 each with pair of dcnihlcd pas. 
Lalerai pas on tarsi lU and IV (latiwc^^iwinult'; 
medial pas on tarsu*} IIP ^btilbrtn, ofi itmsc 1V 

plumose (Fig. 21 A) 

Carpileson tarsi I ;iiid II long. i)ti larsi III ;iiul 
IV 6 long, \ccessory process on each ol' claws 
and wiih 3-4 miniilc tmcs (Fig, 21 A ). 

Female. Idiosonia 205-260 lonii. In dorsal aspect 
SlOJjlar to male- VcnTra! platen Al. IM and <jP 
fijsed; and AP laterally separated by wedges 
offilriaied itiiejpjmSW^Igf ZJl^.'With 5 pairs of 
pg^ 2 .aiaienorpytsipb^Q^^ acteliDr to GO. 
fS) S!l tong. Genifal «c1i^te^ ^jft 2 pairs of sgsf. 
Ovip^Sftor w ith 5 pairs ofgeijiwl spines, 7-11 
fang atld ending with 5-6 tines (Fig. 21C). 
Length:width ratio ofgnaihosiMn.i 1.23. Rostrum 
shorter than gnathosomal base (Fig. 21 Dj. On 
im^^s IV. lalcral pas tkuened and peCsifittlte 
medial pas sctifomi tFig. 21F). 
Tritonymph Idiosonia 204 long. OmAtnerttatioil 
ol' dorsal plates foveatc rather Uian reticulaie 
tf ig. 21G). Posterior margin of AD truncate, 
endiiig immediately posterior to line of muscle 
scars. OC and PD smaller than in adults. Ventral 
plates separated; Gl' iind AP not fused ( Fig. 2 1 H). 
Posterior margni of AI-. con\c.\. At and Pt each 
With pair of adjunct setae. tiP with 2 pairs oFpgs 
jUid Ipairof.sgs.Nuinbei oTselae of trochanter to 
a^aoflegs Landlj: 1,2, 4, 5, 5; of legs III andlV 
i J,433*^d 0, U2, 3. 5. Telofemora t-IV viilh 
3/1, 3/1,3/1, and 2/0 seiae. respeciive!>. 

h'ariQtions. Varieties of characters ui adults: 
length of idiosonia. 9: 205-260 (10), length of 
idic&oma, i^: 205-223 i^i xiMmbvr ol adjunct 
setae on either of AE: 0). 1 (36), 2 (2); 

numher of adjuncl .setae on PF: 1 (?8K 2 (2); 
number of pgs on either side. oF <_jO 9 : 5(20); 
nutnbcr of pus plus basilar sclac in cilliet half, 6 

^1 tU, (7), 8+1 (Uj.'^^l (I); number of 
«eta« ^rfJeg segments 2 to Sj: 



4- 



4/tp4J 



5(40> 



let IV 



lev III 



1U:.MARK.S. Rhomhognathus lericujus closely 
resembles R, reticulifei: In both specresthePfcFis 
reticulate, the ds-l arc ver\' long. AR and PF each 
have 1 pair of adjunct setae, the anal cone is of 
iu>rnid Ihfe gnftthosoitia is short, ihe leg 
vh«etObl3Ky is J^ftw 5iiaila^c,t|}e distajly wi4e^ed 
iapcx of the claws beats ftyt small tbie^ Djf- 
Icivnoes arc: k. tmtnlus has' nmore promin^l 



rctiinxlation, its PDis wider, the telotenvira are 
shorter. ih<; ido&mowill aiid IV ei»ch liave J/I 
dorsal/VCTWats€tafr>'aiiSthatJi^ tihds 
on ihc apex of the claws. TritdByaflphs ol ^\ 
/L'nr/i!n\ liiivea riiveiUe PO, wherWS'lbBffn i /i". 
rt'n'rn/ijtT is reticulated; the OPofA teriOiluSV^ 
larger than in R. n'ncul{Ji't\ 

Both species are eaj^ily sep^atod ft'om siindai 
sized Indo-Wcsl Pacific :spccies on tbc basis of 
the enlarged ds-I. 

KhunibD^iiatbu.s vaiidipes sp.nov. 
F I V MOLOSY. For IB sndng 6^A*i^ LsftoVl^ J^. 

MATI;RIAL, IIOLuTVPL. i (MIQK i'tvoM Biirricr 
JlecC 18^16.46'S, 147=22.88^. M>nuidon Rcct dead 

om, PAiwrYPes. « fOw^owx^oife^ 

above. V, 1 tt-iton>Tnph^UAlDf;/99),odl£^ 
abovT. OTrrr:R MATERTXI.: 1 tritonymph (MTQK tinwt 

Barrier Reef L'ndine Reef oft'Cajv Trihulatioii, giowtli on 
cwals ut hrn, 16 November 1997; aill. J.t'. OiUi. (IBi, 
collection data as abcfy^. 

DliSCRlPFION. Female. Iduv.om.i u s : 
long, holotypc 327 long. 217 wide Dot sal iila'c% 
separated lioni each other hy wide areas ol 
stci&^d integumtmi (Fig. 22Aj. AD 97 lotig^ Ifii 
^dft anterior mar^m biroadly rtnmded, postentir 
margin ovate. Pair of gland pores in lateral 
margin. Fine \\ ith muscle scars at approximately 
0.".V)C slender. 7> and MS long, 2S wide: Fueriil 
margin with 2 conieae and pore canaliculus, Ihc 
larttw closer to poMerior th^n lo anterior gljind 
pore. OC witli 2 corneac, th& pofiierior one 
f?tibdividcd. PD 122 long. Intcgunwttt Wifhirtpair 
of '^li^htK raised co.siae with delicate pores; 
median poiiion of plate w tih faint reticulate orna- 
nicmalinii Fan of gland poies in posletolaieriil 
Vpm^rSy th^ latter sUgJni.l> extending ^(^ond 
Ttscdim moigb) of^E^. An^ ^lei^feS of i^rml 
si/e, Tiot extended by anal valves. Setae ds-l IS 
long; shginly w ider and twice Tbe length of Out 
posterior setae: ds-I insetled at 0.50 relative to 
length of AU. Setae ds-2 and ds-S on OC, Ihe 
postenor opeat 0.64^.68. Setae ds-4 and ds-5 on 
l^DimAmot pmtMt 0,05, posterior sete ^^t 0.47- 
Adahal setae on aiial (Tlate. 

Vcniral phitcs AF!, PFl and GP fused to a ventral 
shield; median portion of AP contiguous lo 
ventral shield, else separated by pair of wide 
4ateral wedges of tahmed iittesiunqit (Tig, 
A& wtthb 2-3 tittuia >i}f ^imist setae; fE-iAlh 2 
wcii^, t^9taD$» from camcf ostofiYc to CO 



196 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 21. Rhombognathus tericulus sp. nov.; A, tip of tarsus IV, ventrolateral, male (dorsomedial fossary seta 
omitted); B, idiosoma, ventral, female; C, ovipositor, lateral, and 3 genital spines (enlarged), female (spines of 
opposite side dotted); D, gnathosoma, ventral, female; E, tarsus I, lateral, female (medial claw and setae 
omitted); F, tip of tarsus IV, ventral, female (dorsal setae omitted); G, idiosoma, dorsal, tritonymph; H, 
idiosoma, ventral, tritonymph. Scale bar = SO^xm. 



1 65 ; GO 67 long. With 5 pairs of pgs, anterior pair 
of pgs at the level of insertion of leg IV. Genital 
sclerites with 2 pairs of sgs. Ovipositor with 10 
pairs of genital spines (Fig. 22C); each spine 
trifid with equal-sized tines. 

Gnathosoma 105 long, 77 wide, 1.36 times 
longer than wide. Gnathosomal base large (Fig. 
22D). Rostrum 27 long, 20 wide, triangular in 
outline (Fig. 22E), much shorter than gnatho- 
somal base. Rostrum with 2 pairs of maxillary 
setae and pair of long lateral rostral setae. Palps 
appressed to and slightly surpassing rostrum. 

Legs stout; their length approximately 0.7 of 
that of idiosoma. Legs III and IV inserted at 0.54 
and 0.68 relative to length of idiosoma. 
Telofemora I and II shorter than these legs' tibiae, 
telofemora III and IV slightly shorter than tibiae. 
Telofemora I and II 1.5 times longer than high 
(Fig. 22F, G); telofemora III and IV 1 . 1 -1 .2 times 



longer than high (Figs 22H, 23A). Leg 
chaetotaxy: leg 1, 1, 2, 7, 6-7, 7, 3; leg II, 1, 3, 7, 7, 
7, 3; leg III, 1, 2, 4, 3, 6, 4; leg IV, 0, 2, 3, 4, 6, 3. 
Telofemora I and II each with 5/2 dorsal/ventral 
setae, telofemora III and IV with 3/1 and 3/0 
dorsal/lateral setae, respectively. Ventral seta on 
each of genua I-IV bristle-like, slightly serrate. 
Both ventral setae on each of tibiae I-IV stout and 
serrate or bipectinate. Tarsus III with 4 dorsal 
setae; the 2 basal setae distinctly separated. Basal 
fossary setae at basis of claw fossa; 2 apical 
fossary setae inserted within fossa area. On tarsi 
III and IV apicalmost (dorsolateral) fossary seta 
near tip of tarsi and resembling medial pas. Tarsi I 
and II each with dorsolateral solenidion, 10 and 
12 long, respectively (Fig. 23B, C); apex with 
pair of doubled pas. Famulus on tarsus I 
papilliform. Tarsi III and IV each with setiform 
medial pas and scaliform pectinate lateral pas. 



RHOMBOGNATHIN AE FROM THE GREAT BARRIER REEF 



197 




FIG. 22. Rhombognarhiis validipes sp. nov., female; A, idiosotna, dorsal; B, idiosouia, ventral; C, ovipositor, 
lateral (spines of opposite side dolled); D. gnaihosoma, lateral; E, gnathosama, ventral; F, leg 1, ventromedial; 

G, leg II, medial: H, leg III, medial. Scale bar = 50^,m. 



Carpites of tarsi 1-IV 6, 7, 10, 10 long. Claws Tritonymph. Idiosoma 267-291 long, 173 wid^. 
smooth. Posterior margin of AD more truncate than in 

female. OC slender as in female; posterior com^ 



198 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 23. Rhombognathus validipes sp. nov., female; A, leg IV, medial, female; B, tarsus I, lateral, female (medial 
claw and setae omitted); C, tarsus II, lateral, female (medial claw and setae omitted); D, tip of tarsus IV, lateral, 
female; E, idiosoma, dorsal, tritonymph; F, idiosoma, ventral, Iritonymph. Scale bar = 50|xm. 



subdivided (Fig. 23E). Ventral plates AE, PE, GP, 
and AP separated. AE with 1-2 pairs of adjunct 
setae; PE each with 2 adjunct setae. GP ovoid 
(Fig. 23F); with 2 pairs of pgs and I pair of sgs. 
Three pairs of genital acetabula adjacent to 
primordial genital slit. Gnathosoma as in male. 
Leg chaetotaxy : leg 1, 1 , 2, 6, 7, 7, 3 ; leg II, 1 , 3, 6, 
7, 7, 3; leg III, 1 , 2, 4, 3, 6, 4; leg IV, 0, 2, 3, 4, 6, 3. 
Telofemora I and II each with 4/2 dorsal/ventral 
setae, telofemora III and IV with 3/1 and 3/0 
dorsal/lateral setae, respectively. Tibiae I-IV each 
with 2 stout ventral setae. 

Variations. Varieties of characters in adults: length 
of idiosoma, female: 315-353 (4); number of 
adjunct setae on either side of AE: 2 (6), 3 (2); 
nimiber of adjunct setae on PE: 1 (1), 2 (7); 
number of pgs on either side of GO, female: 5 (8); 
number of setae of leg segments 2, 4 and 5, and 
number of dorsal/ventral setae of telofemora I 
and II and dorsal/lateral setae of telofemora III 
and IV: 



segment 


leg I 


ieg ri 


leg III 


leg IV 


T 


2(8) 


3(8) 


2(8) 


2(8) 


3 


5/2(8) 


5/2(8) 


3/1(7) 


3/0(8) 


4 


6(1), 7(7) 


7(8) 


3(7) 


4(8) 


5 


7(8) 


7(8) 


6(7) 


6(8) 



REMARKS. Adults and juveniles of Rhom- 
bognathus validipes are characterised by the two 
pairs of setae on the PD, whereas the other 
species from the Great Barrier Reef have a single 
pair of setae. 

In the rhombognathine fauna presently known 
from northeastern Australia, the two species 
Rhombognathus lathridius and R. validipes have 
smooth claws. R. validipes is larger than the 
psammobiont R. lathridius; its gnathosoma is 
wider and the adanal setae do not arise from small 
pedestals as in R. lathridius. 

In dorsal and ventral aspect of the idiosoma and 
the outline of the gnathosoma and the claws, R. 
validipes resembles R. leurodactylus Krantz, 
1976 and R. robustus Bartsch, 1977, both are 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



199 



recorded from the eastern Pacific coast (Krantz, 
1 976; Bartsch, 1 977). In contrast to the two latter 
species, the OC of/?, validipes are more slender. 
Other characters which separate R. validipes 
from /?. leiirodactyliis are: the number of setae of 
the telofemora I-IV - 5/2, 5/2, 3/1, 3/0 in R. 
validipes, but 3/2, 3/2. 2/1, 2/1 in R. 
leurodactylus\ the number of perigenital setae in 
females, 5 pairs in R. validipes^ no more than 3 
pairs in R, leurodactylus. Tibiae I-IV of R. 
robustus bear 6, 6, 5, 5 setae, those oiR. validipes 
7, 7, 6, 6 setae. 

KEY TO ADULT RHOMBOGNATHINES OF 
THE GREAT BARRIER REEF 

I. Gnalhosoma completely hidden beneath AD. OC wider 
than long; without seta. PD with 3 pairs of setae. Venter 
with small plates and large areas of striated integument 
(Fig. IB) Isohactrus poiKipensis 

Gnathosoma extending beyond anterior margin of 
idiosoma. OC longer than wide; with 2 setae. PD with 
1-2 pairs of setae. Ventral AE and PE fused to ventral 
shield Rhombogfiothus 

KEY TO ADULT RHOMBOGNATHUS 

1 . Plates AD, OC and PD separate 2 

All dorsal plates flised. Claws with more than 20 lines 
scutu/afiis 

2. Anterior margin of AD evenly rounded (Fig. 2A). Anal 

plate without pedestals 3 

AD with minule fi-onlal process. Adanal setae on small 
pede.stals (Fig. 7A); gnathosoma 1.5 times longer than 
wide; telofemora I-IV with 5/2, 5/2, 3/1, 3/1 dorsal/ 
ventral setae. Claws smooth lathridius 

3. Telofemora MV with 4-5/1-2, 4-5/1-2. 3/0-1. 3/0-1 dorsal/ 

ventral setae 4 

Telofemora 1-IV with 2/1, 2/1. 2/0, 2/0 dorsal/ventral 
setae cyrtonotus 

4. Claws widened and provided with more than 1 tines (Figs 

15H. 19E) 5 

Claws smooth or with accessory process, the latter with 
l-8tines(Figs5H, I7E) 6 

5. Claws with 12-13 tines. Setae ds-1 longer than following 

setae(Fig. 1 8A); female with 7-9 pairs ofpgs( Fig. 191): 

male with 1 1-1 5 pairs of pgs. - semiuoiatus 

Claws with 18-20 tines. Setae ds-l hardly longer than 
following setae (Fig. I4A); female with 5 pairs of pgs 
(Fig. 15F):malcwilh 10-14pairsofpgs . . . papuensis 

6. Length of ds- 1 not exceeding twice length of following 

setae(Figs4A,9A) 7 

Setae ds-l 3-4 times longer than following setae (Figs 
16A. 20A): anal valves not extending beyond anal 
sclerites; dorsal plates reticulate 10 

7. TelofenioralandlIeachwith4/l dorsal/ventralsetae. . 8 

Telofemora I and 11 each with 5/2 dorsal/venlra! setae . 9 

8. Length:heighl ratio of telofemora 1.9-2.0; anal valves 

extending beyond small anal sclerites (Fig. 4C) 
dclicatulus 

Telofemora slender. Iength;heighl ratio 2.5-2.9; anal cone 
of normal shape with anal valves and anal sclerites 
similar in size (Fig, IIC) longipes 



9. OC 1 .6 times longcrthan wide; two corneae equal in shape; 

PD with single pair of setae (Fig. 9A); claws with 
accessor)' process (Fig. I OA) levigatus 

OC slender, more than 2.5 times longer than wide; 
posterior cornea subdivided; PD with 2 pairs of setae 
{Fig.22A);ciawssmooth(Fig.23R) ..... validipes 

10. PD with iaree-sized, faint reticulum, its meshes 8-IO(xm 
long (Fig. T6A); telofemora I-IV with 4/2, 4/2, 3/0, 3/0 
dorsal/ventral setae; claws with 7-8 minute lines (Figs 
17B, B) reliculifer 

PD with conspicuous and dense reticulation (Fig. 20A); 
telofemora I-IV with 4/1. 4/2, 3/1, 3/1 dorsal/venlra 1 
setae; claws with 3-4 minute tines (Fig. 21 A, F) 

tericulus 

DISCUSSION 

In the Great Barrier Reef area, 1 ! species of 
Rhomhognathus and 1 of Isobactrus were found. 
Future collections will certainly result in records 
of more species. The number of Rhonibognafhus 
species is similar to that known from south- 
western Australia, viz. 10 species (Bartsch, 
1993), and from the boreal and warm temperate 
northwestern Pacific, 1 1 species from Hokkaido 
(Abe, 1996) and 8 species from the Hong Kong 
area (Bartsch, 1992). The number of species of 
Isobactnis recorded from the north- and 
southwestem Pacific area is small. Isohactrus 
ponapensis is the first representative of this genus 
from the shores of Australia. The low number of 
records of Isohactrus may partly be due to the 
lack of collections from adequate habitats. 

Species of Rhombognalhus recorded from various 
regions of the Pacific and the tropical Indian 
Ocean are summarised in Table I. The know- 
ledge of the rhombognathine fauna in the Pacific 
and Indo-West Pacific region still is poor, and 
several of the published records are based on 
sporadic and short-time sampling acti\'ities with 
halacarids being just one of the accessory taxa. 
Accordingly, the data summarised in Table 1 far 
from completely mirror the number and dis- 
tribution of the genus Rhomhognathus in these 
geographic regions. The status of Rhomho- 
gnathus denticulatus Sokolov, 1952 is not clear; 
the absence of a bipeclinate seta on tibia III may 
be an evidence of close relationship to the genus 
Isohactrus. 

Noteworthy is the low number of wide-spread 
species. R. leurodactylus obviously is an 
amphi-Pacific boreal species. The species was 
taken on the shores of Oregon/US (Krantz, 1 976) 
and all around Hokkaido/Japan, here both on the 
coastlines influenced by subarctic water currents 
(Okliotsk Sea and Western North Pacific) and by 
the warm (subtropical) Kuroshio current (Japan 
Sea) (Abe, 1996). R. sinensis inhabits a wide 



200 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Species of Rhombognathus and their records from the eastern and western Pacific and the tropical 
Indian Ocean. 1 ^ North America; II = Hawaii; III = Galapagos; IV South America; V =^ Northern Japan and Sea 
of Japan; VI = Southern China; VII = Philippines, Papua, New Guinea, Guam, New Caledonia; VIII = 
Northeastern Australia; IX =New Zealand; X = Antarctic Pacific; XI = Western Australia; XII = Tropical Indian 
Ocean; * = taxonomic status not clear; x? ^ record in need of re-identification. 



Species 


Geographical Area 


References 


I 


II 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


XII 


acfe/Zewsij Newell, 1984 




















X 






Newell, 1984 


am/j/gHiw Newell, 1984 




















X 






Newell, 1984 


oreKarzwi Bartsch, 1992 












X 














Bartsch, 1992 


atuy Abe, 1 990 










X 
















Abe, 1990, 1996 


bisciitatus Bartsch, 1993 






















X 




Bartsch, 1993 


caudiculus Bartsch, 1983 














X 












Bartsch, 1983 


ceZ)M«5 Bartsch, 1983 














X 












Bartsch, 1983 


compressus Abe, 1 996 










X 
















Abe, 1996 


cyrtonotus sp. nov. 
















X 










present paper 


(itirw/ni Newell, 1984 








X 


















Newell, 1984 


delicatulus sp. nov. 
















X 










present paper 


denticuiatits Sokolov, 1952* 










X 
















Sokolov. 1952 


dictyotus Bartsch, 1 992 












X 














Bartsch, 1992 


dissociatus Abe, 1990 










X 
















Abe, 1990, 1996 


ellipticus Bartsch, 1977 






X 




















Bartsch, 1977 


e/famm Newell, 1984 








X 


















Newell, 1984 


ezoensis Abe, 1990 










X 
















Abe, 1990, 1996 


/e//f/5 Newell, 1984 








X 


















Newell, 1984 


foveolatus Bartsch, 1993 






















X 




Bartsch, 1993 


/rac/z/j Bartsch, 1979 


















X 








Bartsch, 1979a 


glaber Bartsch, 1989 




X 






















Bartsch, 1989a 


guamensis Bartsch, 1989 














X 












Bartsch, 1989a 


heterosetosus Bartsch, 1977 






X 




















Bartsch, 1977 


hirtellus Bartsch, 1992 












X 














Bartsch, 1992 


incertus Abe, 1996 










X 
















Abe, 1996 


msw/ar/s Bartsch, 1989 




X 






















Bartsch, 1989a 


lacunosus Bartsch, 1979 


















X 








Bartsch, 1979a 


latens Bartsch, 1993 






















X 




Bartsch, 1993 


lateralis Newell, 1984 








X 


















Newell, 1984 


/{itfiridizis sp. nov» 
















X 










present paper 


latibulus Bartsch, 1993 






















X 




Bartsch, 1993 


lepidus Bartsch, 1993 






















X 




Bartsch, 1993 


leurodactylus Krantz, 1976 


X 








X 
















Krantz, 1976; 
Abe, 1996 


levigatus sp. nov. 
















X 










present paper 


longipes sp. nov. 
















X 










present paper 


longiscutatus Bartsch, 1977 






X 




















Bartsch, 1977 


longisetus Bartsch, 1 999 














X 












Bartsch, 1999 


lubricellus Bartsch, 1989 




X 






















Bartsch, 1989a 


marginalis Bartsch, 1993 






















X 




Bartsch, 1993 


medialis Abe, 1996 










X 
















Abe, 1996 


mukisetosus 'Newell, 1984 








X 


















Newell, 1984 


neotenus Abe, 1996 










X 
















Abe, 1996 


neptmellus Bartsch, 1992 












X 














Bartsch, 1992 



RHOMBOGNATHTNAE FROM THE GREAT BARRIER REEF 



201 



TABLE \.{cont.) 



Species 


Geographical Area 


References 


I 


IT 


III 


IV 


V 


VI 


VII 


VIII 


IX 


X 


XI 


XII 


novaezelcmdicus Bartsch, 1985 


















X 








Bartsch, 1985 


oblongus Bartsch, 1989 














X 












Bartsch, 1989a 


pacificus Newell, 1 984 








X 


















Newell, 1984 


papuensis Bartsch, 1 989 














X 


X 










Bartsch, 1989a, 
present paper 


placidus Bartsch, 1 993 






















X 




Bartsch. 1993 


plumijer Irouessart, looy 








X 


















JNewell, 1984 


psammopnilus Bartsch, 1993 






















X 




Bartsch. 1993 


reticulatus KranXz, 1976 


X 
























Rrantz, 1976 


reticuiifer sp. nov. 
















X 










present paper 


ro^MSfwi Bartsch, 1977 






X 




















Bartscn, 19// 


scutulatus Bartsch, 1 983 














X 


X 






X 


X 


Bartsch, 1983, 
1993; 

Chatterjee, 1995; 
present paper 


semiarmaTiis Bartsch, 1983 














X 












Bartsch, 1983 


seminotatus sp. nov. 
















X 










present paper 


semireliculatiis Borisch, 1977 






X 




















Bartsch, 1977 


setelhis Bartsch, 1992 












X 














Bartsch, 1992 


Bartsch, 1983 














X 












Bartsch, 1983 


similis Bartsch, 1977 






X 


















X? 


Bartsch, 1977; 
Chatterjee, 1995 


smen^w Bartsch, 1990 










X 


X 














Bartsch, 1990 


sinensoideus Bartsch, 1 992 












X 














Bartsch, 1992 


tenuiformis Abe, 1996 










X 
















Abe, 1996 


tericulus sp. nov. 
















X 










present paper 


terminalis Sokolov, 1952 










X 
















Sokolov, 1952 


teurinus Abe, 1996 










X 
















Abe, 1996 


thalassinus Bartsch, 1993 






















X 




Bartsch, 1993 


validipes sp. nov. 
















X 










present paper 


ventralis Newell, 1984 








X 


















Newell, 1984 


verrucosus Bartsch, 1992 












X 














Bartsch, 1992 



range along the coast of Asia, from Hokkaido to 
Hong Kong (Abe, 1996; Bartsch, 1992). i?. scutii- 
latus is an Indo-West Pacific species; records are 
from the Philippines, eastern and western coast of 
India, Western Australia (Bartsch, 1983, 1993; 
Chatterjee, 1995) and, now, from eastern 
Australia. From the Indian Ocean, Chatterjee 
(1995) published a record of R. similis Bartsch, 
1977, a species also known from the Galapagos 
Islands (Bartsch, 1 977); because of differences in 
the arrangement of the perigenital setae in 
females, the eastern Pacific and Indian Ocean 
specimens may belong to difi^erent species. R. 
papuensis is known from Papua New Guinea 
(Bartsch, 1989a) and eastern Australia. Though 
often not identical species, there are close sim- 
ilarities between the rhombognathines of 



northeastern Austraha and the Philippines, e.g. R. 
lathridiiis/R. ccaidiculus and 7?. cyrtonotus/R. cehuus. 

On the basis of data on the geography and 
ecology of rhombognathines, Bartsch (1982) 
concluded that within Isobactrus many of the 
species have a high tendency to tolerate 
environmental changes but a low speciation rate. 
In contrast, within Rhombognathus there seems 
to be a high evolutionary potential. Rather than 
tolerate a wide range in the environmental param- 
eters, species genetically diversify, and when 
being brought into a new habitat, exposed to 
changing hydrographic or climatic challenges or 
biological interactions, new species evolve in the 
local scene. The large numbers of Rhombo- 
gnathus may be the result of adaptations to the 
numerous niches in the Great Barrier Reef area. 



202 



MEMOIRS OF THE QUEENSLAND MUSEUM 



In the material from the Great Barrier Reef 
Marine Park area at hand, a few individuals could 
not be identified, they demonstrated characters of 
two species, namely R. delicatulus and R. 
tericiiliis. Are these specimens extreme variants, 
separate species, or hybrids? Nothing is reported 
on genetic barriers, or hybridisation, between 
halacarid species sharing a habitat, in the course 
of the author's studies in the northern Atlantic, 
thousands of rhombognathines have been 
examined microscopically. The variation of 
characters is low, apart from a few anomalies, 
namely deformations, intersexes which showed 
characters of both males and females, or adults 
with unilaterally a leg form and setation equalling 
that of juveniles. Hybrid-like forms, with 
characters of two species, were not found. The 
rhombognathine fauna of the northern Atlantic 
includes the genera Jsohactms, Metarhombo- 
gfiatlms, Rhombognathides and Rhotnbogriathus. 
From the Pacific and Indian Ocean only Iso- 
bactnis and Rhombogmitliiis are known. The 
genus Rhomhogfiathiis is characterised by a large 
number of ditTerenl fornis; there are species with 
a slender, elongate idiosoma, others are short and 
flattened; some species are small and cryptic, 
others very large; species have short legs with 
wide segments or the telofemora and tibiae are 
elongate and the legs almost as long as the 
idiosoma; the numbers of setae of the legs vary 
considerably. Beside the wide range of inter- 
specific differences, the genus is characterised by 
a high intraspecitlc variability. In contrast to 
Rhombognafhus, Isobacfrus demonstrates a 
considerable uniformity. The shape of the 
idiosoma and legs is rather similar in all species 
and even interspecific dirterence in the setation is 
small within the faunas of the northern and 
southern oceans, respectively. 

ACKNOWLEDGEMENTS 

I thank Dr J.C. Otto for collecting, sorting and 
providing the halacarid mites described in this 
paper. 

LITERATURE CITED 

ABE, H, 1990. Three new species of the genus 
Rhomhogmilhus ( Acari, Halacaridae) from Japan. 
Zoological Science 7: 517-535. 
1996. Rhombognathine mites (Acari : Halacaridae) 
from Hokkaido. Northern Japan. Publications of 
the Seto Marine Biological Laboratory 37: 
63-166. 

1998. I^ombognalhine mites. Taxonomy, phylogeny, 
and biogeography. (Hokkaido University Press: 
Sapporo). 



BARTSCH, I. 1975a. Ein Beitrag zum System der 
Rhombognathinae (Halacaridae, Acari). Zur 
Morphologic der Tarsalregion und des Ovi- 
positors. Zoologischer Anzeigcr 194: 193-200. 

1975b. Ein Beitrag zur Rhombognathinen-Kauna 
(Halacaridae, Acari) der F^retagnekiiste. 
Acarologia 17: 53-80. 

1977. Interstitielle Fauna von Galapagos. XX. Hala- 
caridae (Acari). Mikrofauna des Meeresbodens 
65: 1-108. 

1979a. Five new species of Halacaridae (Acari) 
from New Zealand. New Zealand Journal of 
Marine (& Freshwater Research 13: 175-185. 

1979b. Halacaridae (Acari) von der Atlantikkiiste 
Nordamerikas. Beschreibung der Arten. 
Mikrofauna des Meeresbodens 79: 1-62. 

1982. Zur Gattung Rhombognathus (Acari, 
Halacaridae), Llbersicht Ciber alle Arten, deren 
Verhreitung und eine Bestimmungstabelle. 
Zoologische Jahrbiicher, Abteilung Systematik, 
Geographic und Biologic derTiere 109: 83-97. 

1983. Zur Halacaridenfauna der Philippinen. 
Beschreibung von Fiinf Arten der Gattung 
Rhombognathus (Acari, Halacaridae). 
Entomologische Mitteilungen aus dem 
Zoologischen Museum Hamburg 7: 399-416. 

1985. New species of Halacaridae (Acari) from 
New Zealand. New Zealand Journal of Zoology 
12: 547-560. 

1986. New species of the genus Rhombognathus 
(Acari, Halacaridae), and a key to Mediterranean 
Rhombognathus. Mesogee 46; 3-7. 

1989a. Rhombognathinae (Halacaridae, Acari) aus 
dem Pazifik. Beschreibung sieben neuer Arten. 
Entomologische Mitteilungen aus dem 
Zoologischen Museum Hamburg 9: 229-246. 

1989b. Rhombognathus auster, a new rhom- 
bognathine mite (Acari, Halacaridae) from 
southern South America. Zoolotrica Scripta 18: 
423-425. 

1990. Halacaridae (Acari) of Hong Kong. Pp. 
661-665. In Morton, B. (ed.) The marine flora 
and fauna of Hong Kong and Southern China II. 
(University Press: Hong Kong). 

1992. Hong Kong rhombognathine mites (Acari : 
Halacaridae). Pp. 251-276. In Morton, B. (ed.) 
The marine Uora and fauna of Hong Kong and 
Southern China III. (University Press: Hong 
Kong). 

1993. Rhombognathine mites (Halacaridae, Acari) 
from Rottnest Island, Western Australia. Pp. 
19-43. In Wells, F.E., Walker, D.L, Kirkman, H. 
& Lethbridge, R. (eds) The marine ilora and 
fauna of Rottnest Island, Western Australia. 
(Western Australian Museum: Perth). 

1999. Rhombognathus longisetiis, a new halacarid 
mite (Rhombognathinae, Halacaroidea, Acari) 
from New Caledonia. Cahiers de Biologic 
marine 40: 15-20. 
BARTSCIL I. & SCHMIDT P. 1979. Zur Verbreitung 
und Okologie einiger Halacaridae (Acari) in den 



RHOMBOGNATHINAE FROM THE GREAT BARRIER REEF 



203 



Sandstranden der Ostsee (Kieler Bucht), der 
Nordsee (Sylt) und des Europaischen Nordmeeres 
(Tromso). Mikrofauna des Meeresbodens 74: 1 -37. 

CHATTERJEE, T. 1995. Record of three species of 
Rhombognathiis (Halacaridae: Acari) from Indian 
Ocean Region. Journal of the Bombay Natural 
History Society 92: 282-286. 

GOSSE, RH. 1 855. Notes on some new or little-known 
marine animals. The Annals and Magazine of 
Natural History (2nd series). 16: 27-36. 

KRANTZ, GW. 1976. Arenicolous Halacaridae from 
the intertidal zone of Schooner Creek, Oregon 
(Acari : Prostigmata). Acarologia 18: 241-258. 

LOHMANN, H. 1889. Die Unterfamilie der 
Halacaridae Murr. und die Meeresmilben der 
Ostsee. Zoologische Jahrbucher, Abteilung 
Systematik, Geographic und Biologic der Tiere 4: 
269-408. 

NEWELL, l.M. 1947. A systematic and ecological 
study of the Halacaridae of eastern North 
America. Bulletin of the Bingham oceanographic 
Collection, New Haven 10: 1-232. 
1984. Antarctic Halacaroidea. Antarctic Research 
Series 40: 1-284. 



SOKOLOV, LL 1952. Vodjanye klesci. II. Halacarae. 
FaunaSSSRS: 1-201. 

SCHULZ, E. 1933. Zur Halacaridenfauna der Kieler 
Bucht. Schriften des naturwissenschaftlichen 
Vereins flir Schleswig Holstein 20: 96-105. 

TROUESSART, E. 1888. Note sur les acariens re- 
cueillis par M. Giard au laboratoire de Wimereux. 
Compte rendu hebdomadaire des seances de 
TAcademie des Sciences 107: 753-755. 
1889. Revue synoptique de la famille des 
Halacaridae. Bulletin scientifique de la France et 
de la Belgique, (serie 3) 20: 225-251. 
1894. Note sur les acariens marins (Halacaridae), 
recoltes par M. Henri Gadeau de Kerville sur le 
littoral du department de la Manche. 
(Juillet-Aout 1893). Bulletin de la Societe des 
amis des sciences naturelles, Rouen 9: 139-175. 

VIETS, K. 1927. Die Halacaridae der Nordsee. 
Zeitschrift fiir wissenschaftliche Zoologie 130: 
83-173. 

1939. Meeresmilben aus der Adria (Halacaridae 
und Hydrachnellae, Acari). Archiv fiir 
Naturgeschichte, Neue Folge 8: 518-550. 



204 



MEMOIRS OF THE QUEENSLAND MUSEUM 



NEW RECORDS OF TRAPEZIID CRABS 
(CRUSTACEA: BRACHYURA: TRAPEZIIDAE) 
FROM THE CORAL SEA AND NORTHERN TASIMAN 
SEA. Memoirs of the Queens/and Museum 45(2): 204 2000:- 
Trapeziid crabs belonging to Tetralia and Trapezia are 
obligate symbionts of reef-building corals, while species of 
Quadrella inhabit ant i path arians, alcyonaceans, gorgonians 
and azooxantheliate corals. The trapeziids inhabiting the 
Coral Sea region were studied by Castro ( 1 997). Examination 
of collections deposited in the Australian Museum, Sydney 
(AM), Queensland Museum, Brisbane (QM) and the Museum 
National d'Histoire Naturelle, Paris (MNHN) has revealed 
the presence of additional species for the region. Two species 
are recorded for the first lime from eastern Australia and two 
from New Caledonia. Five species of Tetralia. twehe of 
Trapezia and three of Quadrella are now known from the 
Coral Sea and northern Tasman Sea. Only one wide-ranging 
Indo-West Pacific species. Trapezia tigrina Eydoux & 
Souleyet, 1848, remains unknown from the region. 

Quadrella coronata Dana, 1852 
For synonymy see Castro, 1999a: 95. 

AMP17448, c?, 9,NewCaledonia, ilotAmadee,30-35m, 
12 Sept. 1929, B. Conseil & G. Bangibout. New record for 
New Caledonia. The only previous Coral Sea record was from 
southwestern Vanuatu (Castro, 1997). The species is known 
from across the Indian Ocean and the western Pacific (Castro, 
1999a). To additional species. Q. maculosa A\cock, 1898 and 
Q. serenei Ga\i\, 1986 (the Vanuatu specimen referred to as Q. 
maculosa by Castro, 1997; see Castro, 1999a: 96), is also 
known from the Coral Sea. 

Tetralia vanninii Galil & Clark, 1988 

For synonymy see Castro, 1999a: 103. 

AMP17232. 2^. Queensland, North Hast Cay, Herald 
(iroup, 17°20'S, 148'^'28'E,6Nov. 1964, D.F. Mc Michael & 
J.C. Yaldwyn; QMW25176, 6,9, Wreck Reef, Porpoise 
Cay,22°17'S, 155°25'E, outer reef slope. 12 m, 10 May 1988, 
J. Short & S. Mullers; AMP39689, f?, 9, Sugarloaf Island, 
Lord Howe Island. 3r30.2'S, l59m3'E, 17m, 27 Dec. 
1 979, N. Coleman. First record for the Coral Sea region. The 
species is found through most ofthelndo-West Pacific region. 

Trapezia formosa Smith. 1869 
For synonymy see Castro, L99S: 178. 

AMP38228. 6, 9, Elizabeth Reef, reef flat near 'Yoshin 
Maru Iwaki' wreck, 29°55.8'S, 159°01.3'E, small coral head, 
J.K. Lowry & R.T. Springthorpe. First record from eastern 
Australian waters. The only previous Coral Sea record was 
from the Chesterfield Islands (Castro, 1997). The species is 
widely distributed across the Indo-West Pacific and eastern 
Pacific regions (Castro. 1998). 



Trapezia lutea Castro, 1997 
Trapezia lutea Castro, 1997: 84-87. figs 2C, D, 3A-C, pis 2C, 5A. 

QMW25186, IS, 9, Queensland, Flinder^s Reef, off 
Cape Morelon. 26°59'S, 153°29'E. RJ.F. Davie & J. Short. 
First record from eastern Australian waters. Previous Coral 
Sea records were from New Caledonia and the Loyalty and 
Chesterfield Islands (Castro, 1997). The species is widely 
distributed through most of the Indo-West Pacific region; first 
recorded from Indonesia by Castro ( 1999b: 52). 

Trapezia punclipes Castro, 1997 
Trapezia piinctipes Castro, 1 997: 87-89, figs 4A-C, pi. 2D. 

MNHN-B26249, ?, New Caledonia, Bale St. Vincent, 
22E25'S, I66H50^E, Aug. 1961, Y. Plessis. The species was 
described from material collected at Lizard Island, northern 
Queensland. This is the first record from New Caledonia. It is 
generally uncommon; known from the Andaman Sea coast of 
Thailand to the Mariana Islands and Fiji (Castro, 1999b). 

Acknowledgements 

I thank Penny Berents (AM), Alain Crosnier (IRD, Paris), 
Peter Davie and John Short (QM) for making this material 
available. 

Literature cited 

CASTRO, P. 1997. Trapeziid crabs (Brachyura: Xanlhoidea: 
Trapeziidae) of New Caledonia, eastern Australia, and the 
Coral Sea. In Richer de Forges, B. (ed.) Le benthos des fonds 
meubles des lagons de Nouvelle-Calcdonie (Sedimentologie, 
Benthos). Etudes et Theses 3: 59-107. 

1998. Systematic status and geographic distribution of Trapezia 
formosa Smith. 1869 (Crustacea. Brachyura, Trapeziidae), a 
symbionl of reef corals. Zoosystema 20(2): 177-181 . 

1999a. Trapeziid crabs {Crustacea, Brachyura, Xanthoidea, 
Trapeziidae) of the Indian Ocean and the Red Sea. Zoosvstema 
21(1): 93-120. 

1999b. Results of the Rumphius Biohisloncal Expedition to 
Ambon ( 1 990). Part 7. The Trapeziidae (Crustacea: Brachyura: 
Xanthoidea) of Indonesia. Zoologische Mededelingen Leiden 
73: 27-61. 

DANA, J.D. 1852. Conspectus Crustaceorum, et. Conspectus of the 

Crustacea of the Exploring Expedition under Capt. Wilkes, 

U.S.N., including the Crustacea Cancroidea Corystoidea. 

Proceedings of the Academy of Natural Sciences of 

Philadelphia 6: 73-86. 
GALIL, B. & CLARK, P.P. 1988. On a collection oi Acropora- 

inhabiting trapeziids (Crustacea Brachyura Xanthoidea) from 

East Africa. Tropical Zoology I: 137-151. 
SMITH. S.I. 1 869. Notes on new or little known species of American 

cancroid Crustacea. Proceedings of the Boston Society of 

Natural History 12: 274-289. 

P. Castro, Biological Sciences Department. California State 
Polytechnic University, Pomona, CA 91768, U.S.A.: 12 May 
2000. 



SPECIES OF DISCOCELIS (PLATYHELMINTIIES; POLYCt^ADlDA) FROM 
QUEENSLAND, WITH QfiSCRIPTlON QF ANEW SPECIES 



Beveridgt 1. 20(H) 06 30 species ol7^/;vcoL-L-7/> (Plat} helriiinthes: \ c ludiJaj frorTi Queens- 
laad»with description of a new species. Memoirs oj the Queensland Museum 45(2); 205-213. 
^sbane. ISSN 007^8835. 

J w o :^pccies of Discocciis are described from int&iS$di waters from northern Qtieenslanii. 
The first species is charaderised by the marginal eyes extending to the posterior region of the 
bod^f the presence of a seminal vesicle, cerebral eyes distributed more or less in anterior and 
mMQtWfi^mi a mate-amnia^ with cOmRlex lobes jtrtlorst^'venttd views^Ttiui ^f^!^ 
difBsfS twny fr^Hi dll «^g6rt6r6 ahd named O. pGtrvilmxcutata ht!Ac Th^seccmd has 
marginal e>'e5 extending to the [eve! of the cerebral organ, cerebral eyes arranged more or 
less in two groups. lacks a seminal vesicle and has prosuiioids in ihe^ wall ofthe male antrum 
as well aji in ilie penis papilla itself. The species is closely re I ait d /^M'V/i/Kato, 1 93 Silt 
is probabi) distinct l->iii is not named owing to the poor description ui / ' p>''\,-''la. The records 
prcscnled suggest that SL'Vcral species oJ DiscoceUs arc present in Austi'alian coastal waters 
and that the distribuiiun of prosiaioids seen in dorso-\ entral views ofthe male antrum 
provides usetui characters for distinguishing ^ieC)6^ Withlt^ gl^Sw O <Pc)/)'C/«)k/^ 
DLsnKelis, ;;cvt spt'cics. taxonomy. 

I. Beverhige, Departmenf of Vetenmry Science, Umversity oJ Melbourne. Parb/Ule 3052, 
MeihauPm* Australia; 24 AprU 



The polyclad ^amly DcseocelidaeLaiiilatv; I9D3 

U a cosmopolitan family of cssentiaily iniertidal 
polyciads characterised by the presence of 
iiKiruinal eyes and .small secretory organs, lemied 
proslatoid.s, associated with tlic male reproductive 
^StefO (f iiubel, 19S3). The family is curretitly 
represented by a single Australian speeieSj 
Discocelis aiis&ah's Uytnsin. 1959, found' Uttdei* 
roeka in die intertidal regit-n close to Sydney 
(Hyman, 1959) and from Wesi I .. South Australia 
CPrudhoe, 1982). Faubel (l')S:v) iransferred /). 
j^mralis^ to thf related Thalamoplatia 
Vaiidlaw;. l$Q4,/(t&lingidslied from Distocetfs 
Elues^bei^ 183$-'fty possessing separate male and 
ferhate gon6p&}«fi» By contrast, Prudhoe (i985j 
considered th^ thiilarfic^lart& ViTdnso^ OfAy 
sub-generic rank. 

Tber piFesi^noe- 6fity a single Atistraliari 
rcpresentalive«ftfi]BfinuIjris probably the result 
oflack of collecting rather than the family being 
pv»or] \ represented in Australian inlertitj^ waters. 
Thivpaper r^)Qrt$ the {jfcssence of tw^addit^Qnal 
ipQws^J>lsso0p To^ville, Queens^ 
land, one of wBichis cieaily specbs. 

METHODS 

Polyciads were coilegt^f) ^ low-tide ironi. 
under rocks on exposed imld-'flats. Kixation 

followed the technique of Newman &. Cannon 
( I995J in which polyciads were placed on filter 
paper in a dish of searw?^^ and wljen fiilly 



esttended, ^(^]^pe!rM<a&Tt^dly placed piEta 
block of frozen fixative, either 4% foBivaldebyd^ 
in sea-water or lormaldehyde-calcium acetafc- 
piopylenc ghcol- propylene phenoxetoi. Following 
fixation, worms were dehydrated in a graded 
series of ethanols, cleared in methyl salicylate 
aad mounted m Canada balsam The median 
i^osterior secfionfi^ of rndividtial polyciads ^Vere 
removed using a scalpel blade, embedded in 
paraffin and serial longuudinal sections, cul al a 
thickness of 7|,im, were stained with Gill's 
haeinatoxylin and cosin. Drawings were made 
usingadi^'ftgtube attached to toOlyttipus BM 
microscope. Measurements are presented in 
millimetres as the range followed by the mean in 
parentheses; 

Ail spec^i^ii^caUQCtedhave beep deposited in 
fhe Quefenslarid Museum XQM). 

Type specunens ot D. a/<Mruli\ from Nc'^n 
South Wales tAu&iraJian Museum W3685) were 
cotnpdtsd VfiSx the new material. 

FOLYCLADIDALana 1884 
ACOTYLEALang, 1884 
DISCOCELIDAE Laidlaw. 190> 

Oiscocetis parvimaculftta sp. luov. 

MATERIAL. HOLOTN PH: Rowc s Ba). To\mis\ iile. (.)ld 
(19''16'S, 146'^49^E), 17.vi.l 997, coll, I. Bexpridge. whole 
mount, unstattied (QM G2 17321); 2 colour slides. 



206 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. Discocelis parvimaculata sp. nov., entire 
polyclad, dorsal view, showing pattern of pigmented 
maculae and extent of marginal eyes (ME arrows). 
Scaie bar = 1mm. 



PARATYPES: 5 entire specimens and fragments of 1 
specimen, whole mounts; 1 set of sections stained with 
haematoxylin and eosin, Rowe's Bay, Townsvilie, Qld, 
coll. I. Beveridge, l.vii.l994, 29.vi.1995 (QM G217322-7, 
serial sections G2 17328). 

DESCRIPTION. Large, oval polyclads; holotype 
non-gravid specimen 13 long, 10 wide; gravid 
specimens, 18-21(19) long, 7- 1 2{ 1 0) wide; dorsal 
surface fawn, darker in centre, covered with 
numerous small brown circular areas of pigment, 
larger brown patches in central regions, becom- 
ing smaller towards periphery (Fig.l); ventral 
surface pale grey; nuchal tentacles absent; 
cerebral organ 0.41 x 0.55 in holotype, 0.39-0.48 
(0.43) X 0.46-0.56(0.51) in paratypes, 2.58 from 
anterior extremity in holotype, 2.94-4.60(3.77 ) in 
paratypes; marginal eyes 3-4 deep, extend to 
posterior quarter of body, number of rows of 




FIG. 2. Discocelis parvimaculata sp. nov., cerebral 
organ, tentacular and cerebral eyes showing variation 
between individual specimens (see Fig. 3). Scale bar 
= 0.1mm. 

ocelli diminishes posteriorly; in most specimens, 
including holotype, eyes reach level of gonopore; 
in some specimens, eyes encircle body; cerebral 
eyes arranged in elongate groups, on either side 
of mid-line, 41-65 ocelli anterior to cerebral 
organ, 5-20 posterior to cerebral organ, anterior 
and posterior groups usually but not invariably 
separated (Figs 2,3); tentacular eyes with 25-40 
ocelli per cluster; rutTled pharynx in mid-body, 
with 10-12 lateral folds, 1 1 in holotype; mouth at 
posterior end of pharynx, 4.9 from posterior end 
in holotype, 7.0-8.1(7.6) in paratypes; single 
gonopore 3.13 from posterior end in holotype, 
5.5-6.8(6.0) in paratypes; antrum masculinum 
volimiinous, folded in both dorso-ventral views 
and sagittal sections; in ventral views (Figs 4,5), 
antrum with prominent anterior lobe containing 
penis papilla and two lateral lobes each partially 
subdivided; in sagittal section (Fig. 6), several 
muscular lobes descend from dorsal surface of 
antrum; antrum with numerous pyriform 
prostatoids opening into lumen; in ventral view, 
prostatoids arranged in subcircular cluster on 
penis papilla and on posterolateral margins; in 
sagittal sections, prostatoids present on all 
pendant processes; no prostatoids present in wall 
of antrum; prostatoids of two histological types; 
most with faintly eosinophic content; prostatoids 



DISCOCELIS SPECIES FROM QUEENSLAND 



207 




FIG. 3. Discoce/is parvimaculata sp. nov., cerebral 
organ, tentacular and cerebral eyes showing variation 
between individual specimens. CE = cerebral eyes; 
CO = cerebral organ; TE = tentacular eyes. Scale bar 
= 0.1mm. 




FIG. 4. Discoce/is parvimaculata sp. nov., gonopore 
and genital complex, ventral view. G = gonopore; L = 
Lang's vesicle; P ^ prostatoids; U ^ uterine duct; VD 
= vas deferens. Scale bar = 0.1mm. 



9°% 




FIG. 5. Discocelis parvimaculata sp. nov., ventral 
aspect showing cerebral organ, eyes, pharynx and 
genital complex. C = cement glands; G ^ gonopore; L 
= Lang's vesicle; M ^ mouth; PH = pharynx; U = 
uterine duct; VD = vas deferens. Scale bar = 0. 1 mm . 



208 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 6. Discoce/is parvimaculafa sp. nov., median sagittal section showing mouth, gonopore and histological 
details of genital ducts. C = cement glands; IN = intestine; L = Lang's vesicle; M ^ mouth; PI = eosinophilic 
prostatoids, P2 = basophilic prostatoids; PH = pharynx; SV = seminal vesicle. Scale bar = 0.1mm. 



at anterior extemity of antrum and on ventral or 
anterior surfaces of pendant processes of penis 
papilla with basophilic content; penis papilla 
fleshy, prominent, in anterior part of antnun; 
ejacLilatory duct simple, straight; prostate absent; 
ejaculatory duct leads to pyriform seminal 
vesicle with thin but highly eosinophilic wall, 
passes venlrally, divides; walls of spermiducal 
bulbs highly muscular; vasa deferentia 
thin-walled, pass anterolaterally from male 
complex, to level of mouth, then divide; posterior 
branches coil posteromedially, uniting posterior 
to Lang's vesicle. No separate female gonopore; 
vagina opens into male antrum immediately 
posterior to common gonopore; vagina with thick 
muscular walls, ciliated lining, cur\^es anteriorly 
to short, horizontal region; uterine canals empty 
into vagina immediately anterior to termination 
of vagina into prominently Y-shaped Lang's 
vesicle; uterine canals extend anteriorly on either 
side of pharynx; cement glands prominent in 
horizontal region of vagina, extend posteriorly 
and laterally into parenchyma, branched distally. 

Discocelis sp. 

(Figs 7-12) 

MATERIAL. Two specimens, Rowe's Bay Townsville, 
Qld, 1 .vii. 1 994, coll. I. Beveridge, whole mount and serial 
sections stained with haematoxylin and eosin (QM 
G2 17329-30, serial sections G2 17331). 

DESCRIPTION. Oval polyclads; gravid speci- 
mens 12-16 long, 5-8 wide; dorsal surface fawn, 
darker in centre, covered with numerous brown 
circular areas of pigment, larger patches in 
central regions, becoming smaller towards per- 
iphery (Fig. 7); ventral surface pale grey; nuchal 
tentacles absent; cerebral organ 0.33-0.45 X 



0.42-0.44, 2.5-4.6 from anterior extremity; 
marginal eyes in rows 3-4 deep, extend around 
anterior quarter of body, reach level of cerebral 
organ; cerebral eyes arranged in elongate groups, 
either side of mid-line, 31-42 ocelli anterior to 
cerebral organ, more or less separate from 4-7 
posterior to cerebral organ (Figs 8,9); tentacular 
eyes with 18-30 ocelli per cluster; ruffled 
pharynx in mid-body, with 10 lateral folds; 
mouth at posterior end of pharynx, 5.2 from 
posterior end; single gonopore 3.4 from posterior 
end; antrimi masculinum voluminous; prominent 
anterior penis papilla, circular in ventral view 
(Figs 10,11), with numerous prostatoids; wall of 
antrum encircling penis papilla bearing single 
row of prostatoids; antrum with 2 laterally directed 
branches on each side, immediately anterior to 
gonopore; anterior pair of lateral branches with 
row of prostatoids along posterior margin; in 
sagittal section (Fig. 12), large muscular penis 
papilla descends from dorsal surface of antrum, 
with numerous pyriform prostatoids; prostatoids 
present in wall of antrum, restricted to anterior 
ventral region; prostatoids with faintly 
eosinophic content; ejaculatory duct simple, 
straight; prostate absent; seminal vesicle absent; 
ejaculatory duct divides into vasa del'erentia 
which pass anterolaterally from male complex, to 
level of pharynx, then re-divide; posterior 
branches coil posteromedially, uniting posterior 
to Lang's vesicle. No separate female gonopore; 
vagina opens into male antrum immediately 
posterior to common gonopore; antrum anterior 
to vaginal opening, prominent, muscular with 
thicker epithelium; vagina with thick muscular 
walls, ciliated lining, ciu^es anteriorly; uterine 
canals empty into vagina anterior to prominent 
dorsal loop; vagina passes ventrally to enter 



DISCOCELIS SPECIES FROM QUEENSLAND 



209 




FIG. 7. Discocelis sp., entire polyclad, dorsal view, 
showing pattern of pigmented maculae and extent of 
marginal eyes (ME arrows). Scale bai' = 1mm. 



Y-shaped Lang's vesicle; uterine canals extend 
anteriorly on either side of pharynx; cement 
glands prominent, extend posteriorly and 
laterally into parenchyma. 

DISCUSSION 

Both species described above belong to the 
family Discocelidae since they possess marginal 
eyes and prostatoids opening into the male 
antrum (Faubel, 1983; Prudhoe, 1985). Generic 
distinctions within the family are not well 
defined, and although both Faubel (1983) and 
Prudhoe ( 1 985) accept the validity of Discocelis^ 
Adenoplana Stummer-Traunfels, 1933 and 
Coronadeua Hyman, 1940, their definitions of 
these genera differ. In addition, Thalamoplana 
Laidlaw, 1904, accepted by Marcus & Marcus 
( 1 966), de Beauchamp ( 1 96 1 ) and Faubel ( 1 983), 
was not accepted as a valid genus by Prudhoe 
(1985). Both species described here differ from 
Coronadena in lacking the 7-11 large prostatic 
organs arranged radially around the male antrum 
in addition to the more numerous small 
prostatoids. Adenoplana was characterised by 
Stummer-Traumfels (1933) as having an 
interpolated prostatic organ. Faubel (1983) by 
contrast interpreted the prostatic organ of 




FIG. 8. Discocelis sp., cerebral organ, tentacular and 
cerebral eyes showing variation between individual 
specimens (see Fig. 9). CE ^ cerebral eyes; CO = 
cerebral organ; TE = tentacular eyes. Scale bar= 0. 1 mm. 



Adenoplana as an ejaculatory duct lined with a 
glandular epithelium. Whatever the precise 
definition of the structures involved may be, 
Adenoplana differs fi"om the species described 
here in possessing distinctly separate gonopores. 

The remaining genera, Discocelis and Tha- 
lamoplana, are distinguishable on the basis of 
gonopores, with the former possessing a single 
gonopore and two gonopores in the latter. 
However, D. australis, which Faubel (1983) 
assigned to Thalamoplana, possesses a single 
gonopore, a feature which was confirmed by 
examination of the type specimens, while D. 
insularis Hyman, 1955 has the male and female 
systems opening at essentially the same point, 
which as Prudhoe (1985) has observed, is 
intermediate between the condition present in the 
type species of the two genera. For the present, 
Faubel's (1983) separation of Discocelis from 
Thalamoplana is accepted but australis is 
considered, following Prudhoe (1985), to be a 




FIG. 9. Discocelis sp., cerebral organ, tentacular and 
cerebral eyes showing variation between individual 
specimens. Scale bar = 0.1mm. 



210 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1 0. Discocelis sp. gonopore and genital complex, 
ventral view. G = gonopore; L ~ Lang's vesicle; P = 
prostatoids; VD = vas deferens. Scale bar = 0. 1 mm. 



member of Discocelis. Both species described 
above are therefore assigned to Discocelis which 
consists of D. australis, D. tigrina (Blanchard, 
1847), D.fiilva Kato, 1944, D. japonica Yeri & 
Kaburaki, 1918 and D. pusilla Kato, 1938. The 
type species, D. lichenoides (Mertens, 1832), is 
considered unrecognisable (Hyman, 1959; 
Faubel, 1983; Prudhoe, 1985) and was treated as 
a species inquirenda by Faubel (1983). 

Within Discocelis, the first species described 
above is immediately distinguishable from all 
congeners on the basis of the extent of the eyes, 
which in other species extend only as far as the 
region of the cerebral organ but in this species 
extend to, or almost to, the posterior end of the 
body. The marginal eyes also extend to the 
posterior part of the body in Adenoplana and 
Coronadena. The species described here differs 
from all congeners except D. australis in 




FIG. 1 1 . Discocelis sp., ventral aspect showing cerebral 
organ, eyes, pharynx and genital complex. C = 
cement glands; G = gonopore; L ^ Lang's vesicle; M 
= mouth; PH = pharynx; U = uterine duct; VD = vas 
deferens. Scale bar = 0. 1 mm. 



possessing a seminal vesicle, though this was 
described as a muscular organ in D. australis by 
Hyman ( 1 959) but has a thin, highly eosinophilic 
wall in the specimens described above. The 
specimens described here differ from D. tigrina 



DISCOCELIS SPECIES FROM QUEENSLAND 



211 




FK-I. 12. Disci'cclis sp., median sagittal section allowing mouth, uonopoie and histological details ot genital 
duels. C = cement glands; IN = intestine; L= Lang's vesicle; P = prosiatoids; VD ^ vas deferens. Scale bar ^ 



andD. mishiitis in b^Ving a male Antrum wtiicti 

foniis five distinctlohesln doi*soventral views. In 
boih of the oTlier species the antrum is rounded, 
based on plale l."^. fiu. 1 of Long ( 18H4) for /.). 
iign'fui and obscr\ations of the type speciincnb in 
the case ofD. aus^raSfff. The morphology of the 
mole lantoa m c(otKbveii!tral view has not 
describra for the remaititng species'. The 
separation of the cerchra! eyes into Iavo elusters 
sepai'ates the species described here from D. 
tigrino^ D. unsfralis and D fulva and the colour 
pauern of the dprsal surface, with numerous 
bf&wn'cirtular aieas separates ^-fepecfes from 

D. Jiilva which lacks a disiinctive pattem (Kato, 
1944). Tltc TWO typc'^ of prostatoids. one with 
eosinophilic content and tlie ulher with bn:^o~ 
philie content may also distinguish this species 
frOJU all congSKtetSyaftbW&llKat (1944, fig. 2) 
iUuste&t^ed types ^prbst^ids^ u) R fyWch 
but did not describe the diffferertccs shown in the 
illustration. The morphological differences noted 
therefore indicate that tl)e described specimens 
represent a new species for which the natne A 
parvimacuicfia is proposed based on the small 
si2e ot^t dbr^Liiid^olar-i^p^d With Dtber 




FIG, T?.SehBI1iaticreprescntalian ofgenilal atriUffiof 
Dtscocelis t^ina^ redrawn from Lang (1 S84). 



speciesiTtwhkirtfiepattpms^onifiedor^l ^tirfece 

ha\ e been adequatofydtescribed. 

The second species described above is 
distinguishable from D, australis and D. 
purvim&culata in lacking a seminal vesicle and 
ftom Ifie tetter -speEiBS in li^ng the marguial 
ey^s restricted to the anterior region of the bodyx 
It differs from /-), (igrioa in ha\ ing the mouth al 
the posterior end of the pharynx rather than in the 
middle and in having Ll>e cerebral eyes divided 
into anWrf*?ratidp09t?riOr groups, in addition, the 
arrangement of the prostatoids in ventral view 
(FigJO) dilTcrS from thai found in D. ii^rinti in 
wliich thev are arranged in a U-.shaped cluster 
around the ainenor half of the peniS- papilla, vvjlli 
two lateral rows extending pOSterfoWy (iMg^ 

ISX4, pi, I V n,y- i )- 

1 lie _specics is therefore most closely related to 
D fulva, D. iitponicif and D- />nsiIl(L all from 
.lapan. The specimens are distinguishable trom 
D Jn/va stA^ this Species- hds rtO dorsal coldUf 
pattern, has rruitt^rous cerebral ^es acranged in a 




212 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 15. Schematic representation of genital atrium of 
Discocelis fulva, redrawn from Kato (1944). 

single elongate group and, according to the 
illustrations of the species, has prostatoids of two 
distinct sizes (Kato, 1944, fig. 2) (Fig. 15). 

D, japouica differs in having 15-16 eyes in 
each posterior cerebral cluster rather than the 4-7 
in the present specimens, and differs in the 
anatomy of the antrum masculinum and distrib- 
ution of prostatoids (Fig. 16). In D. japonic a ^ 
there are a nimrber of projections into the antrum 
apart from the penis papilla, while in the current 
specimens only the penis papilla projects into the 
antrum. In addition, inD. /a/70A?/r(:/, a particularly 
elongate projection, lying dorsal to the vagina 
bears numerous prostatoids on both surfaces (Kato, 
1937, fig. 2), while in the present specimens, the 
region of the antrum anterior to the vaginal 
opening is devoid of projections and prostatoids. 
Unfortunately, no ventral views of the antrum of 
D. japonica have been published. Finally, there 
are prostatoids in the ventral wall of the antrum in 
the current species and these are lacking in D. 
japonica. 

The species described here is most similar to D. 
pusilla in colour pattern, having eyes restricted to 
the anterior part of the body, mouth at the 
posterior end of the pharynx and cerebral eyes 
divided into anterior and posterior clusters with 
only one or two ocelli in the posterior clusters 
(Kato, 1 938). The genital atrium is also similar in 
that there is, according to the illustration of the 
species (Kato, 1938, fig. 3) a large penis papilla 
projecting into the antrum masculinum (Fig. 14), 
although Kato (1938) vStated in the description 
that there were many muscular villus-like 
projections, as in D. japonica. 
Furthermore, there are no 
prostatoids in the posterior region 
of the antrum. The most obvious 
differences between the present 
specimens and D. pusilla are that 
there appear to be very few pros- 
tatoids in the antrum of D. pusilla 
and that prostatoids do not occur in 
the ventral wall of its antrum. 



However, Kato's ( 1 938) specimens of D. pusilla, 
were evidently immature as he describes the 
prostatoids as rudimentary and Lang's vesicle as 
being represented merely by a mass of nuclei. As 
a conseqence, the number and distribution of 
prostatoids may not have been reliably 
determined in D. pusilla. The current specimens 
may therefore be D. pusilla or may represent a 
new species. However, since only two specimens 
are available and since D. pusilla has been 
inadequately described, no new name is proposed 
for them. 

The descriptions presented here indicate that 
Discocelis is represented in Australia by several 
species rather than the single species, D. 
australis, currently known (Hyman, 1959). 
While one of the two additional species found can 
unequivocally be identified as new, limitations in 
the descriptions of existing species prevent a 
definitive name being applied to the second 
species. 

The descriptions presented above suggest that 
in addition to the distribution of marginal eyes, 
the occurrence of cerebral eyes in a single band or 
two groups, and the presence of a seminal vesicle, 
the distribution of prostatoids within the antrum 
masculinum as seen in ventral views of cleared 
specimens provide useful taxonomic characters. 
In D. tigrina, the prostatoids are arranged in an 
arc anterior to the gonopore (Lang, 1884), in D. 
parvimaculata, the prostatoids are arranged in a 
cluster in the anterior lobe of the antrum and 
along the postero-lateral margins while in the 
un-named species the prostatoids occur 
throughout the penis papilla and are present 
along the posterior margin of one pair of lateral 
diverticula within the male antrum. The type 
specimens of D. australis were examined but 
they are now very dark and the distribution of 
prostatoids cannot be determined. In the 
remaining species, this character has not been 
investigated, but current observations suggest 
that it might provide additional features for the 
separation of species within the genus Discocelis. 




FIG. 16. Schematic representation of genital atrium of Discocelis 
japonica, redrawn from Kato (1944). 



DISCOCEUS SPECIES FROM QUEENSLAND 



213 




FIG, 17* Schematic representation of genital atrium of Discocelis austrctlis, redrawn fixmi Hyman (1 959). 



AdKNOWLEDOEMENTS 

DrL.R.G Cannon of the Queensland Museum 
is thanked for comments on a draft of this 
manuscript 

LITERATURE CITED 

de BEAUCHAMP, P. 196L Classe des Turbellaries. 
Twbellari9 QSitr^huir^ 1831), In Grassfi, P>P. 
(ed.) Traits de Zbolo^e 4(1): 35-212 (Nfasson: 
Paris). 

FAiFBEL, A. 1983. The Polycladida, Turbcllaria, 
Proposal and establishment of a new system. Part 

I, The Acotylea. Mitteilungen aus dem 
haraburgischen zoologischcn Museum und 
InstitutSO: 17-121. 
HYMAN, L. 1959. Some Australian polyclads 
(Turbeilaria). Records of the Australian Museum 
25: M7. 

KATO, K. 1937. Pol>clads collected in Idu, Japan, 
Japanese Journal of Zoolog\ 7; 21 1-232. 
1938. Polyclads from Amakusa, Southern Japan^ 

Japanese Journal of Zoology 7: 559-576, 
1944. Polycladida of Japan. Journal tif the 
Sigenkagaku Kenkyusyo 1; 257-318. 



LANCi, A. 1884. Die Polycladen (Seeplanarien) des 
Golfes von Neapel und der angren/enden 
MfciVNahschniUe. Hine Monographic. Fauna und 
Flora des Golfes von Neapel, Leipzig 11: i-xi, 
1-688, 

MARCUS, E. & MARCUS, E. 1966. Systematische 
Uebersicht der Polycladen. Zoologische Beitrage 

12: 319-343. 

NEWMAN, L.J & CANNON, L.R.G 1995. The 
importance of the fixation of colour, pattern and 
form in tropical Pseudocerotidae 
(Platyhelm'inthes, Polycladida). Hydrobrologia 

305: 141-143. 

PRUDHOB, S. 1982. Pohclad lurbellarians from the 
southern coasts of Australia. Records of the South 
Australian Museum IS: 361-384. 

1$&S, A monograph on polycla^ turbellariat 
(Trustees of the British Museum (Natural 
History): London). 

STUMMER-TRAUNFELS, R von. 1933. Polycladida 
(continued). Pp. 3485-3596. In Bronn, H,G (ed.) 
Klassen und Ordnungen des Tier-Reichs (IV). 
(Vermes) (Leipzig). 



NOTOPLANA DUBIA (SCHMARDA) (PLATYHELMINTHES: POLYCLADIDA) FROM 

QUEENSLAND 



1. BEVERIDGE AND T.H. CRIBB 

Beveridge, T. & Cribb, T.H. 2000 06 30. Notoplana dubia (Schmarda) (Plalyhelminthes: 
Poiycladida) from Queensland. Memoirs of the Queensland Museum 45(2): 215-220. 
Brisbane. ISSN 0079-8835. 

The intertidal polyclad Notoplana dubia (Schmarda) (Platyhelminthcs) is reported for the 
first time from Australian coastal waters in Queensland. The significance of morphological 
differences observed between the Queensland specimens and the original descriptions are 
discussed as well as relationships with congeners in Australia and southeast Asia. □ 
Poiycladida. Acotyleu, Notoplana dubia, new record. 

1. Beveridge. Department of Veterinary Science, University of Melbourne, Parkville 3052. 
Melbourne. Australia; T.H. Cribb, Department of Microbiology & Parasitology, UniversiH' 
of Queensland St Lucia 4072. Australia: 28 April 1999. 



Species of the polyclad genus Notoplana 
Laidlaw, 1903 occur commonly in the intertidal 
zone in southern Australia (Prudhoe, 1 98 1 , 1 982). 
The commonest species, N. australis (Schniarda, 
1859), has a wide distribution, ranging from 
South Australia to Sydney (Pmdhoe, 1 98 1 , 1 982) 
and occurs also in New Zealand. Of the remain- 
ing known species, N. longiducta Hyman, 1959 
and A^. longisaccata Hyman, 1959 have been 
described from the Sydney region, while A^. 
distincta Prudhoe, 1982 and N, longicnmiena 
Prudhoe, 1982 occur in South Australia (Hyman, 
1959; Prudhoe, 1981, 1982). No species have 
been reported from northern Australian whalers 
although several members of the genus occur in 
New Britain and southeast Asia (Prudhoe, 1985). 

Faubel (1 983) subdivided Notoplana based on 
the presence or absence of a penis stylet and 
transferred N. longiducta and A^. longisaccata to 
a new^ genus, Notocotnplana Faubel, 1983, 
leaving A', australis, N. distincta and TV. 
longicrumena in Notoplana. Prudhoe (1985) 
adopted a less formal approach, subdividing the 
genus into four groups to facilitate identification, 
the subdivision being based on the presence or 
absence of a penis stylet, penis papilla and penis 
sheath. Prudhoe's (1985) groups A and B 
correspond with FaubeFs (1983) definition of 
Notoplana. Therefore, based on FaubeFs (1983) 
definition, there are currently three species of 
Notoplana in southern Australia, all occurring 
south of the latitude of Sydney. 

This paper reports the finding of a species of 
Notoplana from northeastern and southeastern 
Queensland and its identification as A^. dubia 
(Schmarda, 1 859), the type species of the genus. 



METHODS 

Polyclads were collected at low-tide from 
imdcr rocks on exposed mud-tlats. Mangroves 
(Avicennia spp.) were the dominant trees on the 
shorelines. Fixation followed the technique of 
Newman & Cannon (1995) in which polyclads 
were placed on filter paper in a dish of sea- water 
and when ftdly extended, the filter paper was 
rapidly placed on a block of frozen fixative, either 
4% fonnaldehyde in sea-water or fonnaldehyde- 
calcium acetate-propylene glycol-propylene 
phenoxetol. Following Fixation, wonns were 
dehydrated in a graded series of ethanols, cleared 
in methyl salicylate and mounted in Canada 
balsam. One polyclad was stained in Mayer's 
haematoxylin prior to dehydrating and clearing. 
The median posterior sections of two polyclads 
were removed using a scalpel blade, embedded in 
paraffin and serial longitudinal sections, cut at a 
thickness of 7^m, were stained with GilFs 
haematoxylin and eosin. Drawings were made 
with a drawing tube attached to an Olympus BH 
microscope. All measurements are in millimetres 
and are presented as the range for 1 specimens. 

All specimens collected have been deposited in 
the Queensland Museum (QM). 

Specimens from Queensland were compared 
with the type specimens of Centrostotnuni 
dubium from the Naturhistorisches Museum, 
Vienna (NMV) and of Notoplana evansi Laidlaw, 
1903, now a junior synonym, from the British 
Museum (Natural History) (BMNH). In addition 
other specimens in BMNH and the Swedish 
Museum of Natural Histoiy, Stockliolm (SMNFI) 
were examined and compared with those from 
Queensland. 



21$ 



MEMOIRS Ot THt QUEENSLAND MUSEUM 



POLYCLADIDA Lang, 1884 
ACOTYUEALaqg, 1884 
LEPT0PtANIE»AB5t«npsoi^ 1857 

^oldtitajft& dttbifl <Sdintard^ .1^9} 

(Figs 1-6) 

C'L'iifnj.siafnn/fi duhmm SchmaidO, il4i9 
Inm^enc tmncaid Schmarda. 1859 
Leptoplana duhia Lang, I S84 
I'f^a^cm^emi&i LatiUaw, 

MATERIAL. Four specimens. Port Denlson, Skro S of 
Bowen, QI4 (20°2^S, 148°irE) coll. I. Beveridge, 
3.vii.l994 IQM G217332-5, saial sections G217343): 2 
specimens, Scarborough, Moreton Bay, Qld, (27°12'S, 
1 53'^7T) coll. T.H. Cribb, 1 5.v. 1 m ( C iZ 1 7336-7); 4 spec- 
imens. Weilinijldn Point. Moreton Bu>; Qld, (27''27'S, 
153^14^1- icolLT.l 1. Cribb. 1 1 x.iiU995.1i3t.l997 
(0217338^1. serial scciions 0217342)- 

DESCRIPTK.JN. Large, oval rH>lyclads; mature 
specimens 22-30 long. 13-16 wide: immature 
specimens lS-20 long, 9-1 1 vvide; dorsal surlace 
brownj darker in centre, ventral surface pale grey; 
taichal tenlBd^diininutivei 3^3^6 fran^aittedoc 
margin: derebral organ 0.54-0.3T5«0.52-D.6!; 
eyes arranged in tWO elongate groups, on either 
side of mid-line, 45-55 anterior to cerebral organ, 
15-18 posterior to cerebral organ; mouth 6.S-8.0 
posterior lo cerebral organ; niffjed pharymt in 
mid-body, witii 10-16 lateral folds; TSlnfe genital 
pore 2.8-3 posterior to mouth; antrum 
tnasculinum elongate, with folds: penis papilla 
elongate, prominent, with dislinelive penis 
sheath; penis stylet prominent, 0.57-0.65 long, 
scleroiised, brown in colour; eja^lfttoty duct 
convplwte4 leads to spherical prostate with 
epffli^rliaT tubes lying parallel lo cjaculatory duct: 
seminal xesicle large, inuseulan convoluted, 
passes dorsally then ventraliy, terminating blind- 
ly; vasa deferentia enter seminal vesicle anterior 
to its extreniily, <;oij post<?rolaierally then turn 
anteriorly to testes. Fettiale geiiital dpeiling 
2.0-2 s from male antrum, 4.6-6.5 from posterior 
end: vagma externa prominent with fhick walls, 
leading 10 elongate, horizoiUal \agina media 
surrounded by masses of cement glands; vagina 
media passes anteriorly almost to level of male 
genital ppemijg, ten?wnating in bli?^d diver- 
ticulnm; ifivertJculmn clearly visible in sections, 
not distinguishable in whole mounts, posterior to 
diverticulum, vagina interna passes dorsally eillier 
as sinuous tube, or. in less mature specimens m a 
coil, then leads posteriorly, uterine tanals ejnp^ 
into vagina interna immediately ^nt£^oV 
termination of vagina in inconspicuous Lang's 
vesicle; no histological differentiation noted 
beiM^eeq va^gjii^ iiitenm and L^fV v^^ide; 




1mm. 



uterine capals ex|^d anteriorly on either side of 

pharynx. 

DISCUSSIOM 

The polyclad described aboA c belongs to 
Notoptana, as delined by Faubel ( 1 ^>S3 ). since it 
lacks a sucker, possesses cerebral and tentacular 
but not marginal eyes, an interpolated prostate 
subdivided into longitudinal chambers and a 
prominent penis stylet. The species belongs lo 
Prudhoe's (1985 ) group A since it possesses both 
a penis stylet and a penis sheath. It most closely 
te^cmbles the type specie, A^, dubia, in 
possessing a large, musciuar, convoluted seminal 
vesicle, an elongate vagina media surrounded by 
cement glands and m CilSlSieaae^ dwimUi 
Lang's vesicle. 

N. dubia has beetr descsribed' on a numbef of 
occasions. The original description by Schmarda 
(1859) is very brief but Stiuniner-Traunfels 
(t93^)fTOvided a comptcdi^sive redescription 



NOTOPLANA DUBIA FROM QUEENSLAND 



217 




FIG. 2. Notoplana dithia, tentacles and cerebral organ, 
c = cerebral organ; e = e>e; t = tentacle. Scale bar = 

leased on the original specimens. The original 

description of Laidlaw's (1903) species, N. 
evansi, was poor, but the species \\ as redescribed 
in some detail by Bock (1913), prior to the name 
being made a synonym of JV. dubia by Stummer- 
Traunifels ("1935). Hence, there is ample 
published data on the anatomy of A', duhia. The 
specimens described aho\e from coastal 
Queensland agree in virtually all respects with 
published descriptions of the anatomy of M 
^iuhiay with Ihe eT^ception of the diverticiilum of 
the vagina .-media present in the Queensland 
specimens, which is not mentioned in any of the 
pubHshed descriptions. The type specimen in 
NMV w as examined (NM V 13345) but consisted 
only of the anterior half of the specimen 
preserved in alcohol; the serial sections of the 
posterior region of the bodv were missing. Tlie 
type of , V. c'n//;,s/fBMNH 1949.13.19.7) consists 
of the anterior part of the body mounted in balsam 
and nine slides of transV^e serial sections. 
Because of the transverse orientation of the 
sections, the features of Hie vagma interna were 
not easy to distinguish, but there was no obvious 
indication of a diverticulum or of a coiled or 
sini^ous ascendimgvagjna. Inth^sp^cimi^ ofiV. 




VlQJ^,Votoptamdli^far^y^ andphai>nx- showing 
positions of male and female genital openings, e = 
eye; g= cement glands; I ^ intestine: ni = mouth; ph =- 
pharynx; t = tentacle; u ^ uterine duct; vd ^ vas 
deferens. Scale bar = 0.5mm. 



218 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4. Notoplana ditbia, female genital system, 
ventral view, g = cement glands; Iv =^ Lang's vesicle; 
u ^ uterine duct; v = vagina. Scale bar = 0.1mm. 

dubia described by Prudhoe ( 1950) from Burma 
(BMNH 1950.10.24.8) however, the divertic- 
ulum of the vagina media, while not particularly 
elongate, was evident and the ascending part of 
the vagina interna was straight. Bock's (1913) 
material consists of one whole mount and 20 
sHdes of serial sections. The vagina media was 
clearly visible only in one specimen and a 
diverticulum was detected. In this specimen, the 
vagina media was widely dilated with sperm 
while the diverticulum contained no spenn and 
had only a small lumen. The lack of prominence 
of the diverticulum may be the reason it was 
apparently overlooked by Bock (1913). Assum- 
ing that Bock's (1913) and Prudhoe's (1950) 
specimens are correctly identified, it appears that 
the vagina media of N. difhia has a prominent, 
anteriorly directed diverticulum. The specimens 
described above are identical with those of A^. 
chibia identified by Bock (1913) and Prudhoe 
(1950). The fonn of the ascending region of the 
vagina interna appears to be variable and may be 




FIG. 5. Notoplana ditbia, male genital system, ventral 
view, p ^ penis; pr prostate; ps = penis stylet; sv = 
seminal vesicle. Scale bar= 0.1mm. 



coiled in immature specimens with the coils 
extending into a sinuous or straight duct as the 
animal matures. The necessity of caution in in- 
terpreting coiled or folded structures in polyclads 
which may be subject to maturation and method 
of fixation has been emphasised by Prudhoe 
(1985) and Cannon & Grygier (1991). 

In spite of these limitations, the current spec- 
imens have been identified as A^. ditbia. However, 
the possibility cannot be excluded that the current 
descriptions of A^. dubia include two species, one 
with and one without an anterior diverticulum to 
the vagina media. 

Other species of Notoplana recorded by Pmdhoe 
( 1 985 ) from the Indo-West Pacific region were A^. 
willeyi Jacubova, 1906 from New Britain, N. 
mortemeni Bock, 1913 and N. parvida Palombi, 
1924 from Borneo and A^. tavoyemis Prudhoe, 
1 950 from Burma. A', morteusem was transferred 
to Pleioplana by Faubel ( 1 983 ) and TV. tavoyensis 
to Notocomplana. 

N. wUleyi was described from Blanche Bay, 
New Britain and is similar to N. dubia in many 
anatomical features, differing in being relatively 
narrower and in possessing a more obvious 
Lang's vesicle (Jacubowa, 1906). Jacubowa 
(1906) also distinguished A^. willeyi from A^. 
dubia (=evansi) based on colour, though there is 
no difference in colour between the two species 



NOTOPLANA DUBJA FROM QUEENSLAND 



219 




FIG. 6. Notoplana diibia, transverse section through genital openings, d = diverticulum; g ^ cement glands; Iv = 
Lang's vesicle; p = penis; pr = prostate; sv = seminal vesicle; l = tentacle; u = uterine duct; v = vagina; vd - vas 
deferens. Scale bar = 0.1mm. 



based on Willey's (1897) notes and those 
described above. She also distinguished the two 
species based on the position of the female 
genitalia, but the precise difference utilised were 
not specified (Jacubowa, 1906). Thus the 
differentiation of N. willeyi from A/, duhia 
remains to be verified. Of particular interest in 
the description of A', willeyi the illustration by 
Jacubowa (1906) (pi. 8, fig.S) of a small anterior 
diverticulum to the vagina media. Her sections 
were apparently slightly oblique making the 
significance of the diverticulum difficult to 
evaluate, but the structures illustrated warrant 
further investigation. 

N. parvula is readily distinguishable from A^. 
dubia since it possesses only a short vagina media 
and has far fewer cerebral eyes (Palombi. 1924, 
pL 2, fig. 15). 

N. dubia is a widespread species occurring off 
the coasts of Burma, Malaysia and Sri Lanka 
(Bock, 1913; Stummer-Traunfels, 1933; Prud- 
hoe, 1950). The present report extends this 
distribution to the east coast of Australia. 

ACKNOWLEDGEMENTS 

Thanks are due to Dr L.R.G. Cannon for com- 
ments on a draft of the manuscript, Dr Cannon 
and Dr R. A. Bray for help with obtaining relevant 
literaliare and to Drs D. Gibson, U. Jondelius and 
H. Stattmann for access to specimens in their 
collections. 

LITERATURE CITED 

BOCK, S. 1913. Studien iiber Polycladcn. Zoologiska 
Bidrag, Uppsala 2: 29-344. 

CANNON, L, R. G. & GRYGIER, M.J. 1991. The 
turbellarian Noloplana comes n.sp. 
(Leptopianidae: Acotylea: Polyciadida) found 
with the intertidal brittlestar Ophiocoma 
scolopendrina (Ophiocomidae: Ophiuroidea) in 
Okinawa, Japan. Galaxea 10: 23-33. 



FAUBEL. A. 1983. The Polyciadida. Turbetlaria. 
Proposal and establishment of a new system. Part 
I. The Acotylea. Mitteilungen aus den 
hamburgischen zoologischen Museum und 
InstitutSO: 17-121. 

HYMAN, L. 1959. Some Australian polyclads 
(Turbellaria). Records of the Australian Museum 
25: 1-17. 

JACUBOWA. L. 1906. Polycladen von Neu- 
Britannien und Neu-Caledonien. Jenaische 
Zeitschrift fur Naturwissenschaft 41:1 13-158. 

LAIDLAW, F. F. 1903. On a collection of Turbellaria 
Polyciadida from the Straits of Malacca (Skeat 
Expedition, 1899-1900). Proceedings of the 
Zoological Society of London 1; 301-318. 

NEWMAN*^ L.J. & CANNON. L.R.G. 1995. The 
importance of the fixation of colour, pattern and 
form in tropical Pseudocerotidae 
(Platyhelminthes, Polyciadida). Hydrobiologia 
305: 141-143. 

PALOMBI, A. 1924. Policladi pelagici. Raccolte 
pianctoniche fatte dalla R. Nave 'Liguria' nel 
viaggio di circonnavigazione del 1903-05. 
Pubblicazioni del Istituto di Studi Superior! 
Pratici e di Perfezionamento in Firenze, Sezione 
di Scienze Fisiche e Naturali 3: 3-25. 

PRUDHOE. S. 1950. On some polyclad turbellarians 
from Bunna. Annals and Magazine of Natural 
History (12)3:41-50. 

1981. Polyclad turbellarians from the southern 
coasts of Australia. Records of the South 
Australian Museum 18:361-384. 

1982. Polyclad flatworms (Phylum Platy- 
helminthes). Pp. 220-227. In Shepherd, S.A. & 
Thomas, I.M. (eds) Marine Invertebrates of 
Southern Australia, Part 1. (Handbook 
Committee of the South Australian Government: 
Adelaide). 

1985. A monograph of the polyclad Turbellaria. 
(Trustees of the British Museum (Natural 
History): London). 
SCHMARDA, L.K. 1859. Ncue wirbellose Thiere 
beobachtet und gesammelt auf einer Reise um die 
Erde 1 853 bis 1 857. Bd.I. Turbellarien, Rotatorien 



220 



MEMOIRS OF THE QUEENSLAND MUSEUM 



und Anneliden. 1. Halfte. (W. Engelmann: 

Leipzig). 

STUMMER-TRAUNFELS, R. von. 1933. Polycladida. 
In Bronn's Klassen und Ordnungen des 
Tierreichs, 4, Abteilung Ic, Lieferung 179, 



3485-3596. (Akademisches Verlagsgesellschaft: 

Leipzig). 

WILLEY, A. 1897. Letters form New Guinea on 
Nautilus and some other organisms. Quarterly 
Journal of Microscopical Science 39: 153-159. 



CIRRIPEDES AS PALAEOECOLOGICAL INDICATORS IN THE TE AUTE 
LITHOFACIES LIMESTONE, NORTH ISLAND, NEW ZEALAND 



JOHN BUCKERIDGE 

Buckeridge, J. 2000 06 30: Cirripedes as paiaeoecological indicators in the Te Aute 
Lithofacies Limestone, North Island, New Zealand. Memoirs of the Queensland Museum 
45(2): 221-225. Brisbane. ISSN 0079-8835. 

The Te Aute lithofacies limestone complex lies on the east coast of New Zealand's North 
Island. It is comprised of a series of uniform to poorly bedded calcarenites and coquinas of 
late Neogene age. This limestone complex was deposited olT-shore, to the east of the current 
land mass, and during the late Tertiary-Quaternary period was structurally accreted to the 
mainland as a series of longitudinal prisms. This paper reviews the current interpretation of 
the depositional environment, which implies a facies change in the limestone from shallow 
water in the west, to deeper water in the east. Primarily utilising barnacle palaeobathymetry, 
it is shown that one of the oldest and most westerly limestones (the mid-Pliocene Titiokura 
Limestone) was deposited in moderately deep water. 

Living barnacles have clearly detlned bathymetric ranges and temperature tolerances. When 
barnacles are abundant in fossil remains, especially of late Cainozoic age, they are often 
useful palaeoenvironmental indicators. The barnacle fauna of the Titiokura Limestone is 
characterised by the presence of the deep water balanomorph Pachylasma. Species of 
Pachylasma are widely distributed in the living shelf fauna, being found in middle to outer 
shelf environments, although on very rare occasions specimens may be found living in 
waters as shallow as 55m. Therefore, on the basis of the abundant and comparatively well 
preserved plates of Pachylasmcu it may be inferred that the Titiokura Limestone accumulated 
in moderately deep, off-shore conditions. 

This interpretation is confounded however, by the presence, in the same horizons, of a 
species of an exclusively intertidal balanomorph, Epopella. Whilst it is apparent that the 
bathymetric ranges of some taxa change through time, all known species of Epopella are 
demonstrably intertidal to uppermost subtidal. They are also characteristic of temperate waters. 
Sedimentological observations suggest that the Titiokura limestone is a mixed thanato- 
coenosis that accumulated in the middle to outer shelf environment. Shallow water elements 
were introduced as components within submarine avalanches and slurry deposits. Intertidal 
temperature regimes were probably similar to those currently existing along the New 
Zealand coastline. □ Te Aute lithofacies limestone, balanomorph cirripedes, Quaternary, 
palaeobathymetiy, palaeotemperatures, submarine avalanches. 

John Buckeridge, Auckland University of Technology, P.O. Box 92006, Auckland, New 
Zealand: 7 July 1998. 



Cirripedes (barnacles) are very much a 
Cainozoic phenomenon. Charles Darwin was so 
taken with their post-Cretaceous abundance that 
he described Tertiary seas as abounding 'with 
species of Balanus to an extent now quite un- 
paralleled in any quarter of the world' (Darwin, 
1 854). The Sessilia, or 'acorn' barnacles, are also 
the ocean's great opportunists, their choice of 
substrate, both animate and inanimate, being 
unparalleled amongst the invertebrates. Many 
sessile barnacles have become sufficiently 
specialised to be host specific (Buckeridge, 1 998). 
Cirripede palaeoecology is a relatively recent 
pursuit (Foster & Buckeridge, 1987), with barnacles 
traditionally interpreted by non-cirripede specia- 
lists as shallow water taxa. However improved 
sampling techniques, in conjunction with deep 



ocean surveys and recent advances in cirripede 
phylogeny and biogeography, show a distribution 
extending well beyond shelf and slope, with taxa 
recorded from depths in excess of 4000m 
(Buckeridge, 1997; 1999). 

What is of great interest however, is the 
paiaeoecological inferences that we can make 
regarding balanomorph barnacles through time. 
The earliest confirmed balanomorphs are known 
from the Palaeocene of the Chatham Islands 
(Buckeridge, 1983; 1993; 1996). These include 
taxa within the genera Bathylasma and 
Pachylasma in what, from sedimentological and 
associated faunal associations, are clearly 
shallow water sediments (Buckeridge, 1 993). No 
deep water sessilians are knovm from the early 
Palaeogene. 



222 



MEMOIRS OF THE QUEENSLAND MUSEUM 




1000 



100 



10 







c 




Tetraclitidae 




; 7 






-MIIMIIII 


L i 1 1 1. 1 i 1 ui ui ui Jimmiim. 


IMIIlll 


65 


35 







millions of years B. P. 





1000 



100 



10 



Chthamalidae 



: 7 WM^^^^^^ 



D 




llllllllllllllllllllliliiiiininiiiiimitiiiii 



65 



35 

millions of years B. P. 



FIG. 1. Schematic representation of preferred environments for 'primitive' balanomorph families in time. 
Vertical scale: estimated water depth (in metres). A, Pachylasmatidae and Bathylasmatidae; B, 
Archaeobalanidae; C, Tetraclitidae; D, Chthamalidae. All taxa are interpreted as arising in shallow water 
environments, with the absence of pachylasmatids and bathylasmatids from shallow water, and chthamalids 
from all except the uppermost intertidal, being a result of competition, particularly from selected 
archaeobalanids and the balanids. The balanids become abundant in the Miocene, with many species being 
characterised by active feeding, high metabolism and early onset of sexual maturity. 



The first deeper water balanomorphs are re- 
corded from the earliest Miocene, Cape Rodney 
Formation in Auckland's Hauraki Gulf 
(Buckeridge, 1 983). There, large numbers of dis- 
articulated plates of Bathylasma aucklaudiciwi 
(Hector, 1888), are found in beds surrounding 
fossil sea mounts. The inferred depth of deposit- 
ion for these beds is in excess of 100m. In the 
Neogene, species of Bathylasma and Pachylasma 
are not found in shallow water facies anywhere, 
with only one living species, Pachylasma 
japonica Hiro, 1933, being found occasionally at 
depths of less than 80m. The deepest water 
balanomorph known is the bathylasmatid 
Tetrachaelasma tasmanicum Buckeridge, 1999, 
which is extremely abundant, both living and as 
'sub-fossir, adjacent to deep sea mounts of the 



South Tasman Rise in depths of 2200-3600m. 
Preliminary investigations of the balanomorph 
fauna at, and adjacent to, these sea mounts show 
distinct zonation of the Sessilia, with the 
balanomorph species Bathylasma aleariim 
(Foster, 1978), and the verrucomorph^///vem/ca 
gibhosa (Hoek, 1883) restricted to a shallower 
zone of 800-2300m. An important characteristic 
of the deep sea barnacle accumulations is that 
they have a very low biodiversity. This is 
certainly the case with the Tetrachaelasma beds 
on the South Tasman Rise; further, the lowest 
horizons of the Cape Rodney Formation contain 
abundant Bathylasma cmcklandicum, but rarely 
any associated macro-fauna except another deep 
water barnacle, Metaverntca recta (Aurivillius, 
1898). Interestingly, stratigraphically higher 



CIRRIPEDES AS PALAEOECOLOGICAL INDICATORS 



223 



fossiliferous horizons in the Cape Rodney Form- 
ation are supplemented by an inllux of shallow 
water elements, introduced as 'turbid slurries'. 

Modem shallow water and intertidal environ- 
ments in New Zealand and Australia are 
dominated by chthamalids, tetraclitids and 
balanids (Fig. 1 ). The first two groups are knowi 
only from shallow water: chthamalids, in par- 
ticular, are confined to the upper littoral zone, 
whilst tetraclitids extend to the middle littoral. 
Providing no evidence of transportation can be 
demonstrated, this restriction makes both of these 
families very useful in defining shallow water 
marine facies. 

THE TE AUTE LIMESTONE COMPLEX 

The Te Aute lithofacies limestone is a dom- 
inant part of the physiography of the central to 
southern east coast of New Zealand's North 
Island (Fig. 2). It is comprised of a series of 
uniform to poorly bedded calcarenites and 
coquinas of late Neogene age. This limestone 
complex was deposited off-shore, to the east of 
the current land mass, and during the late 
Pliocene-Quaternary was structurally accreted to 
the mainland as a series of longitudinal prisms. 

The term 'Te Aute Limestone' was first used in 
1887 by James Hector in his summary of 
Alexander McKay's geological report for the 
region (Hector, 1887; MacKay, 1887). At that 
time, Te Aute was the name of an area char- 
acterised by extensive limestone outcrops, south 
of Hastings in central Hawke's Bay. The term Te 
Aute lithofacies limestone' was coined by Beu 
(1995) and reflects a greater complexit}' in the 
limestones of the region than was perhaps 
perceived by earlier authors, such as Kimmia 
(1971) and Beu et al. (1980). Beu (1995) estab- 
lished that the Te Aute lithofacies limestone was 
deposited Irom Tongaporutiian to early Nuku- 
maruan age (late Miocene to earliest Pleistocene). 

Of particular note is the type and nature of the 
biota comprising the Te Aute lithofacies 
limestone. Unlike tropical limestones, which are 
characterised by high levels of lime mud 
(Campbell et al., 1988), the temperate conditions 
under which the Te Aute lithofacies limestone 
was deposited has resulted in a much higher 
proportion of skeletal material. Of particular 
interest here is the remarkable dominance of 
barnacle remains in many horizons. Kamp et al. 
(1988) recognise the bamacle content to be 
approximately 13-22% of total rock volume, with 
the highest percentage characteristic of the Scinde 
Island Formation (outcropping near Napier). The 



most impressive 'cirripede dominated lithologies', 
however, occur in horizons of the Castlepoint 
Formation, also of Nukumaruan age, which out- 
crop to the southeast of the Te Aute limestones. In 
a coarse coquina outcropping at Castlepoint, I 
have estimated balanomorph content (primarily 
Fosterella tubulatus (Withers, 1 924)), to be close 
to 50%. 

PALAEOENVIRONMENT OF THE 
TE AUTE LITHOFACIES LIMESTONE 

Although barnacle limestones are not common, 
sediment dominated by shells of the barnacles 
Notohalanus vestitiis (Darwin, 1854) and 
Austronwgahalanus decor us decor us (Darwin, 
1 854), is currently accumulating at depths of 30m 
in the Hauraki Gulf, Auckland, inner to middle 
shelf deposits, dominated by species of the 
balanomorph Fosterella, were a feature of cooler 
southern seas in the Pleistocene (Buckeridge, 
1983). However, with the extinction of Fosterella 
in the early Holocene, no bamacle species has 
moved to dominate the shelf en\ ironmenl to the 
same degree. 

Better sampling and recording of the living 
fauna in recent years has shown that balano- 
morph barnacles can be significant contributors 
to sediments forming at depths of 2000m or more 
(e.g. extensive beds of Tetrachaelasma spp. are 
knovm from the Southern Ocean ofi* Madagascar, 
Chile, Cape Horn and Tasmania). In some of 
these locations, the barnacle shells are found to 
comprise more than 90% of the calcitic remains 
(Newman & Ross, 1971; Buckeridge, 1999). 

The balanomorph shells preserved in the Tc 
Aute lithofacies limestone are comprised of calcile 
rather than aragonite (as for many bivalve mol- 
luscs). The greater chemical stability ofcalcitehas 
contributed to the dominance of balanomorphs in 
many limestone horizons, although this is not the 
prime factor for their abundance. The environment 
suited barnacles, especially the now extinct 
Fosterella tubulatus that is so common in Nuku- 
maruan lithologies. Apart from molluscs, the 
other relatively abundant groups preserved are 
bryozoans, which possess both aragonitic and 
calcitic skeletons. .According to Kamp et al. ( 1 988), 
brv'ozoans never reach the abundance of the 
barnacles in the Te Aute lithofacies limestones. 

PALAEOTEMPERATURES. Barnacles are 
generally of value in interpreting palaeo- 
temperatures through association, e.g. species of 
the reef coral symbionts Hexacreusia and 
Creusia are characteristic of tropical seas. The 



224 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2. East coast of New Zealand's North Island, 
showing locaHties mentioned in text and distribution 
of Pliocene 'Te Aute Uthofacies limestones' and 
Mesozoic Basement. TL = summit outcrop of the 
Titiokura Limestone, AL = Kahuranaki outcrop of 
the Awatapa Limestone. (Modified from Kamp et al., 
1988). 

tetraclitid, Austrobalanus imperator (Darwin, 
1854), is currently known from shallow tropical 
and subtropical seas off central and northeast 
Australia. It has also been recorded from the 
Oligocene of the South Island, a period when sea 
temperatures were at least as warm as the present. 
Not siuprisingly, it is not found in present New 
Zealand waters. Species of Epopella, another 
tetraclitid, are found throughout New Zealand 
and Southern Australia. Epopella sp. is also 
present in the Awapapa Limestone, outcropping 
on the slopes of Mount Kahuranaki (Fig. 2). 
Chthamalids are usefiil indicators of temperature, 
with the endemic New Zealand species 
Chamaesipho brunnea (Moore, 1944) restricted 



to warm, temperate intertidal environments. Beu 
(1995) and Milliman (1974) suggest that the Te 
Aute lithofacies limestone was deposited in 
'shallow subtidal waters'. If this is accepted, the 
absence of intertidal species (with thick, solid 
shells like C. brunnea), indicates that 
temperatures were likely to have been cool 
temperate. This is consistent wdth the molluscan 
evidence of Beu (1995). 

PALAEOBATHYMETRY. Current interpretation 
of the depositional environment (Kamp et a!., 
1988; Beu, 1995), implies a facies change in the 
limestone deposits, from shallow water in the 
west, to gradual deepening conditions in an 
easterly direction. Further to the east again, 
shallower conditions developed, with the uplift 
of the East Coast Highlands. It is perceived that a 
broad NE-SW trending, open seaway, with strong 
tidal currents (the 'Ruataniwha Strait') charac- 
terised the waters between the two land masses. 

In Kamp et al. (1988) and Beu (1995) it is 
proposed that iarge carbonate sand dune-forms' 
colonised by a barnacle-bivalve-bryozoan 
assemblage, formed at the margins of the 
Rualaniwha Strait, and in a moderately high 
energy 'tidal' enviromnent, migrated to deeper 
water (i.e. mid-channel). However, Beu (loc. cit.) 
also refers to Milliman ( 1 974) who states that the 
porous nature, low resistance to abrasion and 
fragility of barnacle plates ensures that they 'do 
not survive transportation from living sites of 
more than a few lens of metres'. Strong cross 
bedding, with tangential foreset beds in tabular 
sets is recognised by Beu (loc. cit ) as confinning 
that the depositional environment was shallow, 
and dominated by strong tidal currents. 

There are some unusual anomalies here, 
particularly in light of the barnacles present in 
some mid-Waipipian limestones in the west and 
central regions. The Titiokura Limestone, out- 
cropping to the northwest, is characterised by a 
mixed assemblage of barnacles, including 
Pachylasma sp., Aiistromegabalanus mio- 
deconis Buckeridge, 1983, and Epopella sp. c.f. 
E. plicata (Gray, 1843). The Awapapa Lime- 
stone, which outcrops in the central region south 
of Hastings, also possesses a mixed barnacle 
assemblage, including^, miodecorus, Notobalanus 
vestitus, Fosterella tubulatus and Epopella sp., 
but lacking Pachylasma. As discussed earlier, 
Pachylasma comprises exclusively deep water 
taxa; further, like Epopella the shell is 
non-porous. If the shell is moderately thick, as is 
the case with large specimens o^ Epopella, it is 



CIRRIPEDES AS PALAEOECOLOGICAL INDICATORS 



225 



likely that transportation, albeit with some 
abrasion, is possible for distances of many 
hundreds of metres. Wliat is even more important 
is that Epopella is found in the same horizons as 

Pachylasma. 

ANALYSIS 

A revised sedimentary regime for the New 
Zealand mid-Pliocene is proposed here. Utilising 
barnacles as indicators of palaeobathymetry, it is 
suggested that the western deposits, such as the 
Tiliokura Limestone, represent a depositional 
environment in water depths of 100m (or more). 
Shallow water sediments, building up on the 
margins of the Ruataniwha Strait, were triggered 
by local overloading, and perhaps micro-seisms, 
to carry intertidal elements, including Epopella, 
as slurries into the deeper water. The process, 
although not unlike traditional turbidites, differs 
primarily in scale, with the distances sediment 
being transported, and the energy involved in that 
process, being at least an order of magnitude less 
than anticipated in 'flysch' deposition. None- 
theless, the mixed thanatocoenosis that the 
Titiokura Limestone represents is consistent with 
deposition from a submarine avalanche, which, 
during the process of its journey, incorporated 
intertidal (Epopella), subtidal {Austromega- 
balanus) and deep water elements {Pachylasma). 

It is also proposed that the Awapapa Lime- 
stone, with a barnacle fauna comprising elements 
from intertidal {Epopella). subtidal (Ausfro- 
megabalanus and Notobalamis) and upper-mid 
shelf (Fosterella) environments, formed in a 
similar manner, but was deposited at depths of 
only 30-40m. Although there aie some similar- 
ities, the Awapapa Limestone almost certainly 
formed in slightly shallower conditions than the 
extraordinary 'Fosterella coquinas' of the 
Castlepoint Formation, outcropping further to 
the south. 

ACKNOWLEDGEMENTS 

I thank Dr Alan Beu, Institute of Geological 
and Nuclear Sciences, Wellington for thoughtful 
discussion at the inception of this study. Mark 
Northover, Learning Technology Centre, 
UNITEC, provided assistance in the production 
of Figure 2. 

LITERATURE CITED 

BEU, A.G 1995. Pliocene limestones and their scallops. 

Institute of Geological and Nuclear Sciences 

Monograph 10: 1-243. 
BEU, A.G, GRANT-TAYLOR, T.L. & de Homibrook, 

N.B. 1980. The Te Aute Limestone facies. 



Poverl>' Bay to Southern Wairarapa. 1:125,000. 
New Zealand Geological Survey Miscellaneous 
Map Series 13 (2 sheets) and Map Notes (36pp.). 
(Department of Scientific and Industrial 
Research: Wellington). 
BUCKERIDGE, J.S. 1983. The fossil barnacles 
(Cirripedia: Thoracica) of New Zealand and 
Australia. New Zealand Geological Survey 
Paleontological Bulletin 50: 1-151. 
1993. Cirripedia and Porifera. In Campbell H.J., et 
al. (ed.) Cretaceous-Cenozoic geology and 
biostratigraphy of the Chatham Islands, New 
Zealand. Institute of Geological and Nuclear 
Sciences, Monograph 2: 1-240. 

1 996. Phylogeny and Blogeography of the Primitive 
Sessilia and a consideration of a Tethyan origin 
for the group. Crustacean Issues 10: 255-267. 

1 997. Cirripedia : Thoracica. New ranges and species 
of Verrucomorpha from the Indian and South- 
west Pacific Oceans. Memoires du Museum 
national d'Histoire naturelle 176: 125-149. 

1998. A new coral inhabiting barnacle, 
Chionelasmus cwsnieri sp. nov. (Cirripedia: 
Balanomorpha) from New Caledonia, Southwest 
Pacific. Zoosystema 20(2): ) 67-1 76. 

1999. A new deep sea barnacle, Tetrachaelasma 
tasmanicum sp. nov. (Cirripedia: Balanomorpha) 
from the South Tasman Rise, South Pacific 
Ocean. New Zealand Journal of Marine and 
Freshwater Research 33: 521-531. 

CAMPBELL, C.S., KEANE, S.L. & HEAD, P.S. 1988. 
Non-tropical carbonate deposits on the modem 
New Zealand shelf Sedimentary Geolog\' 60: 
71-94. 

DARWIN, C. 1854. A monograph on the sub-class 
Cirripedia, with figures of all species. The Ba- 
lanidae, the Verrucidae. (Ray Societ>': London). 

FOSTER, B.A. & BUCKERIDGE, .I.S. 1987. Barnacle 
Palaeontolog>'. Crustacean Issues 5: 43-62. 

HECTOR, .1. 1887. Progress report, 1874-76. Geol- 
ogical Survey of New Zealand. Reports of 
Geological Expeditions during 1874-76. 9: i-xiii. 

KAMP, P.J.J., HARMSEN, FJ., NELSON, C.S. & 
BOYLE, S.F 1988. Barnacle-dominated lime- 
stone with giant cross-beds in a non-tropical, tide 
swept. Pliocene forearc seaway, Hawke's Bay, 
New Zealand. Sedimentary Geolog>' 60: 1 73-1 95. 

KJNGMA, .r.T. ]97L Geology of the Te Aute Sub- 
division New Zealand Geological Survey Bulletin 70. 

McICW, A. 1887. Report on country bet\veen Cape 
Kidnappers and Cape Turnagain. Geological 
Survey of New Zealand. Reports of Geological 
Expeditions during 1874-76 9: 43-53. 

MILLllVLAN, J.D. 1974. Recent sedimentary carbon- 
ates. Part 1 - marine carbonates. (Springer- Verlag: 
Berlin). 

NEWMAN, W.A. & ROSS, A. 1971. Antarctic 
Cirripedia. American Geophysical Union 
Antarctic Research Series 14: 1-257. 



226 



MEMOIRS OF THE QUEENSLAND MUSEUM 



FASTKRN RANGl I MTNSION FOR MOHETHlA 
RVFICAUOA COURTESY (JF A CAT STOMACH 
^fymoMf the Quetimlanii Aiii^euM 4^(2) 226. 20()0:~ 
[dk!ntiffcasm:df mamma] ion n:mums Lrum prcduior pcliet^i 
and scats has provided vfiltiablc infotmatioR an tbe 
distribiitioits of manly spede^'S^fth O^^TVIdoTitSfled ^4 
species of nfttive mammals thai Had iioi bc^ ptofiijiSsS 
rccoi dcd in ihe Flinders Ranges ol'South Australia nbm 
pctlei deposits foLUid in caves. Mure rcecntU . onalysis) iif 
Larnixitrc sluk Wd ro J nunihk^t of signiricanl range extensions 
oi* species ihai arc rare lu-.d or dilTicuIl la dcJietL using 
coiivenlioiiaJ iaiinA cctuisittw icLhriitiucs (fbtri^XMDplC' MeoK 
&. Triggs. K^^J? and iclvTcnecs within ). 

Such Icchniqucb lvj\c' ruii been .r. successlul U>r repiiles. 
wliich uMiaHv lawk ihc hurd bi»dv p^irw recpjirt'd Inr 
ideniitlcarKMi IV-llowiogpassagc through (he guioraprcdalor. 
Siomacii conicdi s of predators, however, oftcr a better chance 
of idenutying reptile remains to species level. Probably Ihe 
beat known result from this type ol* sampling tvW ififl 
redisvovm of the Adelaide oymty bliis-tm&p iWifm 
adetatd,.'tim\ Irom the ittomacn w liT efii5ilam orcwn sniike 
{Psciuiofiatu ^AmsOinlWfArmsiroiig&teld, 1:902). This 
mw rcpiiris J vonsiderubic range extension for Motvthia 
riif'icaudii n)ilo\ving ilic identitication of tbi^species from the 
siurnjL h of a tcial cai iruin Dtamaminalak^sNsitona] Parte 

in rar-wesleni Qiiecnslaml 

1 he skink remains (hindbody. pelvic regiOnandtflll basej 
were Ibirnd in tlie sloruaeh ol a male cat shot by the Park 
Ranger during the da} al the Diamantina (iales (2V4^'S. 
14 l''08't)onthc UihJune 1994, I he remains ucre tdenritK-d 
on the conibinalion oTsi/e and colouration, parlieukiri) th*: 
width, inlcnsily and position of the pale dorsolaieral stripes, 

piamwt^ Kfver'cut^attlie Diamantfni QfeocfS'Vftryiiftlinar 
Ih cbepr^ffewed hahiiai af M rtffkuudit describedby Ctiggft# 

\mi). 

Cogger (1992) lists the disiribuiion ul AI ru/tcamlo as the 
continenl, Irum \\ c -^I 1 1| the 
Noftbcrlt Tcrritoiy. Queensland Itorder exiendui- \>- the 
Western Australian coast and north tollic Morihcrri l en uon 
eoasl. More recentU. A/ ruhtuiiiiu was recorded in ihe 
spinileA duncllelds ol ihe nnrihern Simpson Dcs.-t i i ?^v4ri'S. 
1_'1.X'^2KT* ) iti western i,»ueensland (t3ownc\ Dickinan. 

and I'roni I awn Hill National Park in far imrihwcMern 
Queeniiturid (I8"35',S. l.>K^35Ti (McKas Ckirkc. 1999). 
No voucher specimens were eollccieti tVoin these sites. A 

1im$dUM&tra^ dt ^e Riverstefgh n>siril sim fTSknvwutheast 

oI* laiwn Hill "National Park) in Mj> 1*^87 (T. Couper. pers. 
comm.), A voucher -Specimen otihis species Irom Riversleigh 
(20kni nonhwesi o\ the honiesteadj was coUQGlpd in 
1 989. but ha.s only recently bi4ui'U«^ge|fwt!lh^ti1)fiaeD&^ 
museum (OH J7I441). 



Mclariand < 1 "-J^-iH ) Tailed to uncover any further records ot* 
this species in an extensive review of hTstcffical inlormaiioii 
on the fduna ol" the Channel Counir\ bUiregion of 
suu^eslem Queensland. Ouf record extends the toiown 
7ang& o£this^l(^>e» by apptioxtmatftlv 2701cm iti:atr 

rthesol^ea^4^m1]|pLawIin] National Park obsiervgtiQnt 

A T^kfr Of cat diBts by Dickman (1996) tbundt^st 
repiiles WW relatively iximilton diejat> items during slimmer 
especially indrierareas.CatstGodtoprcy mostly on noeiumat 

species and dragons, w hich reflects their nocturnal hunting 
habits, i hme\ er, undci tlu- tin- enndilions cxpcnenced in the 
Diamantina re\:;i(Mi durini:; l'-'^4. cats were eommonh seen 
huniiny on mild a\ uiier da\ l"hij t) pe ul behax iour wa> not 
ohsen ed in the sub^equenl > ears ot the suid> w hen conditions 
improved and staple prev ripeeits more abundant (R. 
Palnriertp*5r$. obs).Thu>j, it is noi surprising that cat ,slumaclis 
•99il?ctipd during ]:994 contiun«d a large number af dmmal 

Acknowledgments 

This Work was part of a lai-ger siud}- on feral i?a(s tlindv:dbj 
1-nvironnienl 'Vustialia. f hunks goes to *Chooic' HoberUOn 
for providing this and other cat ston>ucli& 

ARMSTRo^ r r a Rrif) i This t^cowMyflfibc Adelaide 
rrtGorR II G RepatMimd«nj9biManspf.Aus4«l|^ 

Books S> unev i_ 

Dlt 'kM W'- C K \' '*'->Ci rn^rview of the impact*, of fer.-il cWi ott 
\u.--irjlijii nuinc r.itinii. (ANCA. Canberrat. 

DOWNEY, FJ. & DtCkMAN. CM. ^<m. Macrn- mi.\ inicmhabirar 
yelfttiOttSllipS ajnong liz^ird- .-f r-jndr,agc dcriL-n m central 
Ausiraito, Pp. 133-138. Ill Liir,nj\ W Aver;., D. leda) 
nprrclolDg) in Atislralta-. .d . lt,:: I'l-^i; plau', fR.tyal 
^^<K>logiofll Society of New Siuili \\ ilcv; MiiMujai. 

McTAKI \M t IX Faun:! ol'ihd ', liLiimel Coui|l^ bi^pflgr^tuc 

ivuHni ioutl". 'Jijtijns[:nid Report \S9 ^fit^lD^^^ 

UcpartinenL vit K.-niinniiviu =jnd Ilcritnge- 

McKAY.I-. atCLARiCE, R.l ( i w. Soniercirtile range exicnsicmsai 
Lawn Hill' Walioniil P;irk, (itilf of CarpcntariH. Queensland, 
M.,-ir':i..i.ii!''.'i V'i \ i: :i-2a, 

ME-.I !n. I' I I l i' /GiS. 13 ^"*7. L%Lpnsion lo ihe rantie nf the 
ca.\terii chi:siniji niouj*, Aitf^t/cJWJ'^ gnKHicttmitUUi. 10 Jen is 
aL^,^|4e«^UUi Wdl««. AU$trfl]tdn lytoAiAktp^ ^1): 

SMI I M \t 1 1' R. rrK'. ii-. <->i' iii[iinmal&. including Snitmns 
.''i . . .■■■.■l.'i ' 1. nuikli I t".oilt.-ni!,r, Miiridae). in owl petleii froin 
i liiidcrs tiajigev bA AusiraLiaJi Wildlilt! Research 4: 159-170- 

Putmer. Deportmeni of ZaethgyandBit/nmefogy, 

Unfyiersfrf of Oueemfaml St Luciit. 4U72: 0.^^ ^i^it, 
Ot-pwlmcnf t>J Veterin^r} -tnafofvy wxi Pafhoio^^^ t^ierat^ 



STEM CASTS OF LEPTOPiJLOELM AISTRALE (WQWy ) WALTON ANDIHB 
RESPONSE THEIR LEAF-CLlSHlON ^ASES. TO COMPRESSION 



tt. TREVOR CUFFOIU? Afctp-lVftpMAEL R. PEMBEBTONT 



<.lltTord. H.I ^ Pemb^rWO, M.R. 2U(lft fl6 3fl: Sicni c-asls Lcpfophiccnffi unsfrulc 
(McCoy) WalioijatrdJI^tespDnseotUieirlear-cushioabascs lo compression. Mw/w'rxo/ the 
gmi9simiJi4its&Nns4S{2y:W-^^jr Brisbane: ISSN 0079-Bh2t5. 

Thedwtribntion patterns ttTthc \\ idlhs o/'ieai'-cushion bases on tii j >i irt":jces of I cpcophhenm 
australe casts itiay be used lo dctemikie vvhether the> >vere eliipi iiul or circular in scciiow 

firiorto their compression. Given the outline $hap^ of a C4$t and iis maximum v\ idlh aivl 
eji^th in section the d^sm&itr oftte oti^inal stem can be j^alciUaicd. O Leptophlaeum. 

H. Trevor CItffbrd QueemM Mmum. PO^X S300. Somh Bri^fxm^mi jWJe^/f. 



C asts and moulds of Lcptophlne^inf nusfraJc 
(McCoy) Walton have been fcporlcd thmi Upper 
ijevbnianand Lower Carboniferous fresh-water 
deposits throughout JLaurasia ( Anderson & 
Anderson, 1985; Cai & Wang, DaMi?on 
IS62, McCoy, IS'74) and r)oiuK\*ana e\ccp! for 
South America and Aniarcliea ( l^d\* ards Sl Berr>, 
■). Theafeeitce of the species frora these two 
continents may merely reilect inadequate 
collecting because in ihose itmes they were 
closely connected with the remainder of Goo* 
dwana (Scotesc & McKcrnm. l^^Of 

The casts derive mainly from the infilling of 
liuHow stems whose central tissues -detsaycd 
lca\ing onl\ a thin sheath of outer cortex and 
icaf^usliion bases. The evidence for the stems of 
Lepfoph/ucum being herbaceous is compelling. 
None of the casts studied taper and none bear the 
scars of sited branches as do those of Lepi- 
dodendron {L'lodLnuiron}: the outlines of the 
leaf-cushion bases, irrespective of the cast 
diaineler, are all of the same shape and meet 
along Iheir margins; in conUasi the leal-cusliion 
bases on larger Lepidodemko)) casts "flre often 
laterally distended and arc more or less separaied 
thereby eonfii mini; that they grew out from stems 
which had experienced secondjry growih- 
Assuming the stems of LepTophlo^nm were 
iier?Jijfi(ittMSfc^*ttay"tei3ompared uuh those of 
preKent-day Agas-e spp. which shortly after 
flowering rapidly decay except for a sheatli of 
encircling leaf-bases. Tliough A<!.itye stems are 
imlially erect and circular in section (Pig. 1 ) the 
4yiag or dead flowering stems soon topple to the 
gyoimd where, tinder the influence of gravity, 
they become elliptical in outline. As a result of 
the slow 4ecay of tiie sheathing' leaf'*ba5)es«'wjU} 



die passage of time, the minor axis of the ellipse 
becomes shorter and Us major axis longer, 

Irtllliing and burial of the hollow stems of 
Lept^i^&mcniStruiere&vS^xiXht Ibrmatidtibf 
intWnslBasl^ <^^i"' ^vhose suriaces are imprinted the 
outlines 6f the leaf-eushiun bases as seen from 
the centre ofilie stem ( fig. 2A). Their outlines str^ 
rhombaidal and ib^ hav&concave surtacen (Fi^ 
2C>. Therefore the Icaf-cmhion impressions oil 

moulds derived 'i [hese ca'.ts arc rhomboidal 
in oulUne with convex surfaces. Casts of whole 
Stem segm^ts -with Setsrils 5f liie stem sitrfee^ 

preserved thereon are rare and readily distin- 
guished Ironi internal casts. Depcndmg upon the 
amount of decay experienced by the stem surface, 
prior to the focni^fixinof the cast, tlie outlines of 
the leaf-cushion feaScs arc more or less distinct 
but their surfaces are always conA cx ( Fig. 2 B.I )) 
Consequently the sitrfaces of the leaJ^ushion 
base£; on moul4£k d^iVtod from such j^dst^^ 
concavB= 

By far the majority of easts deri>e from 
decaying stems vvlfich are ahgncd parallel to the 
bedding plaj'ves of the host sediments. In section 
these easts are elliptical in ouilinfi with their 
major axes disposed parallel to the bedding 
planes and so ai e presunied lo have formed from 
axes that had toppled over at the site of growth or 
had t)eeti transported to a swamj> before infilling- 
Only two casts w vtli eyiindiical sections have 
been noted. One had eroded frOlU its investing 
matrix and was collected ftom a surficial deposit 
(Queensiand. Mines l^paxUn^Qt L1526); the 
other was erect atfd pteSfirvcd in situ (Morris, 



228 MEMOIR^ 51? TflE^UEEkSt ATiJD Mti SEt/M 




FlGl . Erect decaying sterrt of Agave ifivipam. 



MATERIAL AND METHODS 

An internal cast of l^cftuphlncnni itustrult' 
whiph i^i ellipticyl in section and has ihc pattern of 
leaf-cushion bas^ preserved over its whole 
surface was the starting point for tliese studies 
(Clitlord, 1996). Assuming, that before burial 
and compression, the cast wits circularin section, 
the Icnglli of its perimeter, was ascertained by 
muhipiying the number of leaves per whorl by 
fh^ w\dtji} qt'tlip least distorted le^&^usl^pu 
t3n fts stJrfece. tJsing this length the diameter t>f 
the uncompressed stem was calculated. The 
result rc\ calcd ihat the length ol thc major axis of 
the elliptical cast and that oflhc diamclcr t*f'(he 
uncompressed circular cast w ere similar lliereby 
GOnfirmmg llial during comprc v^u)!! the width of 
the cast Imd PQt expanded in the plane at right 
angles t6 the directbrt of ihe apj^licd pressure. 
Therefore, it can hp assumed that when 
compressed, buried casts suffer little or no lateral 
expansion. Because the shape and width of 
leaf-cusliiun bases in whorl are sjanilai^ their 
distiibaiian ^dimcf a circular stem may fee 
expressed as equal arcs on (he perimeter of the 
circle which results Irom sectioning the stem at 
right angles to its length. Likewise, the 
distribution of leaf-cushion bas^s around the 
perimeter of the ellipse dcfivedbV C^tolffcssion 
of a circle may ^sq be represetit6d% a series of 



FIG. 2. Diagrajnmalic \ iesv ol" a decavmg stem ol 
LejUifphiDciim aiisfrijlc and two caslb derived 
t!>er*^from^ A, It-ansvfise section of stem; B, 
l<iOgitojSj|jU|ls&Ctionot sten\; C, internal cast; D,stem 
«avity^ teaf-cushian; It, leaf-trace;; ost, 
tJtiter steni tissue. 

equal-length arcs. The procedure for determining 
the co-urdinatcs which detlne the margins of the 
individual leai-cushion bases around the 
perimeter of an ellipse is complex. However, 
with the a<J\ cni mbolio inanipidators such as 
Maple, iimflieinaiieal [iu-.blenis like those 
encountered in this paper are easily s^Ked. 
M^pte V Relc^e 4 wajs chuspn {o furnish \bc^^ 
t^^vAts as described' Appeitdix I . Wher^s 
during the compression of a circle into an ellipse 
the length of its perimeter is conserved, the length 
of tile major axis ofthe ellipse alvsays exceeds the 
diameter ol the parent eircle (Fig. 3), 

To explore the responses of leaf-ctishion base 
widths to compression, a circuTar east t)f uftit 
diameter (for practical purposes the unit 
employed was 1 00mm) with 32 leaves per whorl 
was taken as a model. This number of leaves was 
chosen both because it is close to that obser\ edon 
lllE cast prev iously studied (C lifford, 1 966) and is 
exactly divisible by four thereby giving a whole 
number per quadrant. TIte theoretical responses 
t)f leaf-cushion bases to compression were 
in\ cstigated for the four uncompressed elliptical 
and one circular cast vvhose sections are 
illustrated inFig= 3. The perimeters of all sectiom 



STEM CASTS OF LEPTOPHLOEUM AUSTRALE 



229 




FIG. 3. Circle (a) and four of the ellipses (b-e) which 
result from its compression. 



are of the same length and each bears the same 
number of evenly distributed equal-sized 
leaf-cushions. The circular cast was subjected to 
four levels of compression (Fig. 4) and the 
elliptical to 3, 2 and 1 level respectively (Fig. 5). 
The dispositions of the leaf-cushion bases around 
the perimeters of the sections of the theoretical 
casts may be calculated as arc lengths but because 
of the irregularities which occur on cast surfaces 
it is customary to measure the widths of their 
leaf-cushion bases in terms of the chord lengths 
joining their margins. Therefore to pennit direct 
comparisons of the amount of change in the 
widths of leaf-cushion bases which occurs during 
compression, chord rather than arc lengths were 
investigated throughout this study. With 
theoretical models chord lengths are calculated 
from the co-ordinates which define the 
boundaries of the leaf-cushion bases, but on casts 
the chord lengths are measured directly with a 
pair of dividers. 

RESULTS 

The relationships between arc and chord 
lengths for two leaf-cushion bases from the same 
whorl about the perimeter of two casts, one of 
which is circular and the other elliptical in section, 
are shown in Table 1 . 

The manner in which the widths of leaf- 
cushion bases, expressed as chords, vary on the 
surfaces of theoretical casts ofLeptophloeum that 
have been subjected to several levels of 
compression are given in Table 2 for an initially 
circular, and in Table 3 for initially elliptical, 
casts. In both Tables the width of the leaf-cushion 
bases is expressed as a fraction of that prior to 
their compression. 




FIG. 4. Circle (a) and four of the ellipses (b-e) which 
result from its compression subject to the condition 
that the major axes of the ellipses and the diameter of 
the circle have of the same length. 



DISCUSSION 

Whereas for a circle the relationship between 
chord and arc lengths is constant for leaf-cushion 
bases arranged around its perimeter, with an 
ellipse this relationship depends on the position 
of the leaf-cushion base on its perimeter (Table 
1). On surfaces of little curx^ature arc and chord 
lengths are similar, but when the curvature is 
considerable arc lengths are conspicuously 
longer than chord lengths. Nonetheless, except 
where the curvature is extreme, the difference 
between the length of the chord and its associated 
arc is insufficient for the former not to stand as a 
surrogate for the latter. Indeed, the greater the 
compression the more closely the chord length 
approximates to the arc length on surfaces of 
minimum compression (Table 1 ). It is clear from 



TABLE 1. Chord lengths expressed as a fraction of 
their associated arc lengths for two nodes on the 
perimeters of a circle and those of four ellipses 
derived therefrom by compression, a-e and 1-8 as in 
Fig. 4. 



Perimeter 


Chord length 


Node 1 


Node 8 


a 


0.99839 


0.99839 


b 


0.99947 


0.99486 


c 


0.99981 


0.98393 


d 


0.99994 


0.96106 


e 


0.99999 


0.95799 



230 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. Relative widths of leaf-cushion bases 
expressed as a fraction of their original widths for 
eight nodes around a quadrant of the perimeters of 
four elliptical casts derived by compression without 
lateral expansion of one initially circular in section, 
b-e and 1-8 as in Fig. 4. 



Node no. 


Length of minor axis of ellipse as a percentage of 
the diameter of the parent circle 


80(b) 


60(c) 


30 (c) 


20 (d) 


1 


1.00 


1. 00 


1.00 


1.00 


2 


0.99 


0.97 


0.96 


0.96 


3 


0.96 


0.92 


0.90 


0.89 


4 


0.92 


0.86 


0.81 


0.78 


5 


0.89 


0.79 


0.71 


0.66 


6 


0.85 


0.71 


0.59 


0.50 


7 


0.82 


0.64 


0.48 


0.35 


8 


0.80 


0.61 


0.41 


0.22 



Tables 2 & 3 that irrespective of whether the cast 
was originally circular or elliptical in section, the 
widths of leaf-cushion bases, respond in a similar, 
though not identical, manner to compression. 

Furthennore, the amount by which successive 
leaf-cushion bases differ in width around the 
perimeter of the cast retlects both its original 
shape and the amount of compression it has 
experienced. If the original cast was circular in 
section a reduction in width by 10% or less is 
experienced only by those leaf-cushion bases 
immediately adjacent to the position of minimum 
curvature on its surface. That is, in proceeding 
from the position of minimum to that of 
maximum curvature on the cast surface there is a 
relatively rapid decrease in leaf-cushion base 
widths but the amount of decrease between 
successive nodes is not constant. Close to the 
positions of greatest and least curvature on the 
cast surface the difference in the widths of 
adjacent leaf-cushion bases is less than for pairs 
of leaves midway between the two (Fig. 6.A). 

In contrast, if prior to its compression the cast 
was elliptical in section, a reduction of 10% or 
less in width is experienced by several, rather 
than a few of the leaf-cushion bases close to the 
position of minimum cur\ ature. Furthennore, in 
progressing from the position of least to greatest 
curvature on the surface of the cast the ditler- 
ences in the widths of successive leaf-cushion 
bases increases consistently (Fig. 6B).Therefore, 
provided they are sufficiently represented, the 
relative widths of successive leaf-cushion bases 
in the same whorl offer a reliable guide to the 



TABLE 3. Relative chord widths of leaf-cushion bases 
expressed as a fraction of their original widths for 
eight leaves disposed around the perimeters of three 
families of ellipses each resulting from the 
compression of an ellipse. n-p,r-s and u as in Fig. 5; 
numbers 1-8 also as labelled in Fig. 5. 



Node 
no. 


Minor axis of initial cast/minor axis of compressed cast 


40/30 
(n) 


40/20 

(0) 


40/10 
fp) 


30/20 
(r) 


30/10 
(s) 


20/10 
(u) 


1 


1.00 


1.00 


1.00 


1.00 


1.00 


1.00 


2 


0.99 


0.99 


0.99 


1.00 


1.00 


1.00 


3 


0.98 


0.97 


0.96 


0.99 


0.99 


1.00 


4 


0.96 


0.93 


0.91 


0.98 


0.97 


0.97 


5 


0.95 


0.87 


0.84 


0.96 


0.94 


0.98 


6 


0.88 


0.78 


0.71 


0.93 


0.88 


0.96 


7 


0.81 


0.65 


0.53 


0.85 


0.75 


0.92 


8 


0.76 


0.52 


0.30 


0.70 


0.45 


0.68 



cross-sectional shape of the cast before its 
compression. 

GENERAL DISCUSSION 

The shape of a cast prior to its compression 
though of interest, is often of less importance 
than a knowledge of the length of the diameter of 
the stem from which it derived. If the leaf- 
cushion pattern on an elliptical cast indicates it 
was originally circular in section, then the lengths 
of its major axis and that of the stem diameter are 
the same. However, if the pattern of leaf-cushion 
bases indicates that prior to its compression the 
cast was eUiptical in section, estimation of the 
diameter of the original axis is more complicated. 
The complexity arises because two processes are 
involved. The first of these is the collapse, under 
the iniluence of gravity, of the original circular 
hollow stem into one elliptical in outline before 
becoming infilled to form a cast; the second is the 
response of that cast to compression. Because the 
girdle of leaf-cushion bases encircling the 
decaying circular stem forms only a narrow band 
of tissue its collapse under the influence of 
gravity results in the stem assuming one of an 
infinite number of elliptical outlines. The length 
of the major axis of this ellipse is fixed prior to its 
being infilled to form a cast (Fig. 1). With one 
exception, the lengths of the major axes do not 
provide a direct estimate of the size of the original 
cast. The exception is when the transecfional area 
of the cast approaches zero in which circum- 
stance the length of the perimeter of the ellipse 
approaches twice that of its major axis. Such 
ribbon-like casts are abundant and derive from 
hollow stems which have collapsed completely 



STEM CASTS OF LEPTOPHLOEUM AUSTRALE 



231 





FIG. 5. Ellipses resulting from the compression of 
three ellipses (b-d of Fig. 4), each subject to the 
constraint that the length of its major axis is not 
affected by the compression. 

or almost so, before being infilled to form a cast. 
The surface of such ribbon-like casts corresponds 
with that of the parent stems and so their leaf- 
cushion bases have convex surfaces (Fig. 2D). 

Wlien the minor axis of the elliptical section is 
much shorter than its major axis, the perimeter of 
the original circular stem can still be estimated 
provided the amount of compression is knowii. 
One estimate of this amount is provided by the 
cast previously reported upon (Clifford, 1996). 
During compression its cross-sectional area was 
reduced to 50% of the original. As this cast is 
typical of all others encountered it has been 
assumed that they too will have experienced a 
similar amount of compression. Therefore the 
lengths of minor axes of casts known to have 
been elliptical in section before compression will 
have been reduced by a half although their major 
axes will not have changed in length. The minor 



axis of the uncompressed cast must be twice that 
of the observed cast, a statement that is confinned 
by the following relationship between the area of 
an ellipse and that of its axes: 

A^-rrab 

where A is the area of the ellipse, a is the length 
of its semi-major axis, b is the length of the 
semi-minor axis. 

Given the lengths of the semi-major and 
semi-minor axes of an ellipse the length of its 
perimeter may be calculated. For high accuracy it 
would be necessary to use elliptical integrals to 
resolve the problem but for the present purposes 
the following approximation is sufficiently 
accurate: 



(a 



+ b + ^JUdT^^) ) 



where P is the perimeter, a and b are defined as 
above. 

It should be noted that to determine the 
perimeter of a cast prior to its compression the 
value of b to be employed is twice that observed 
on the compressed cast. The above discussion has 
assumed that the shape of the cast previous to its 
compression was detennined from a study of the 
dimensions of the leaf-cushion bases on its 
surface. However, even if the leaf-cushion 
pattern is incomplete or absent an estimate of the 
diameter of the original cast is still possible 
assuming that during compression about half its 
volume has been lost. Accepting such loss the 
shape of a cast which is elliptical in section prior 
to compression may be determined by comparing 
the length of twice its minor axis with that of hs 
major axis. If the two lengths are similar the 
original cast will have been circular in outhne; if 
double the length of the minor axis is less than 
that of the major axis the original cast will have 
been elliptical in outline. These results stem from 
the simple relationship between the area of an 
ellipse (of which the circle is a special example) 
and the length of its two axes. Should twice the 
length of the minor axis exceed the length of the 
major axis it would follow that the cast had lost 
less than 50% of its original volume, a situation as 
yet not encountered. The procedure for 
detennining the diameter of a stem whose cast 
was elliptical before its compression will now be 
demonstrated using data from a specimen in the 
collection of the Australian Museum (F6449). 
The outline of the cast as seen in transection is 
shown in Fig.7 together with some points on the 
perimeter of the ellipse calculated from its 
maximum and minimum width. The close 



232 



MEMOIRS OF THE QUEENSLAND MUSEUM 




NODE NUMBER 



NODE NUMBER 



FIG. 6. A, The lengths of the leaf-cushion base widths, expressed as a fraction of their uncompressed value, for 16 
leaves arranged around the semi-perimeters of the four ellipses (b-e) illustrated in Fig. 4; B, The lengths of 
leaf-cushion base widths expressed as a fraction of their uncompressed value for ! 6 leaves arranged around the 
semi-perimeters of the ellipses n-p,r-s and u illustrated in Fig. 5; node numbers 1-8 and their mirror images 
(italics) as in Figs 4 and 5 . 



agreement between the outline of the cast (Fig. 7, 
a) and the superimposed points supports the view- 
that the cast arose by deformation of one which 
was circular in section. 

Doubling the length of the minor axis of the 
ellipse (Fig. 7, a), without altering the length of its 
major axis results in a new ellipse (Fig. 7, b) 
which may be taken to represent the transectional 
outline of the cast prior to its compression. How- 
ever, as it is postulated that the uncompressed cast 
arose by the infilling of a circular but hollow stem 
which had collapsed into an ellipse under the 
intluence of gravity, the length of the perimeter of 
the uncompressed cast and that of the stem will be 
the same. The width of the persistent outer stem 
tissues within which the uncompressed cast is 
fonned may be neglected for it is small compared 
with the diameter of the cast The perimeter of the 
uncompressed cast may be calculated from the 
formula cited above, and using the result, the 
diameter determined of a circle with the same 
length perimeter. This circle (Fig. 7, c) may be 
taken to represent the outline of the original stem. 
It should be noted that whereas the transectional 
area of the compressed cast is only 40% that of 
the stem it is 50% that of the uncompressed cast. 
The difference between these two numbers arises 
because during the deformation of a circle into an 
ellipse the length of the perimeter is unchanged 



but the enclosed area is reduced. Had cast F6449 
resulted fi'om the direct compression of a speci- 
men which was circular in section it would have 
lost c. 80% of its volume. In this circumstance the 
length of the diameter of the uncompressed cast 
and that of the major axis of the compressed 
elliptical cast would have been the same. If such a 
large loss of volume is acceptable, the width of 
the uncompressed cast would be intermediate 
betw een the length of the major axis of the cast 
and that of the diameter of a circle derived on the 
assumption that the hollow stem had collapsed 
into an ellipse before infilling and then losing 
50% of its volume. At present only one cast is 
known whose surface pattern is sufficiently well 
preser\'ed to allow its precompression, tran- 
sectional area to be estimated with confidence. 
Therefore until further data are available it is 
necessary to accept that 50% compression of 
Leptophloeiim axes is the norm, at least in eastern 
Australia, and to use this value together with the 
maximum and minimum widths of casts to 
calculate their precompression diameters . 

ACKNOWLEDGEMENTS 

It is a pleasure to thank the Director of the 
Australian Museum and Dr Sue Palfrey of the 
Queensland Mines Department for providing 
hospitality and access to the collections in their 



STEM CASTS OF LEPTOPHLOEUM AUSTRALE 



233 



jlD: 










-til 6c 



FIG. 7. Outline of an internal cast of Leptophloeum 
atistrale (Australian Museum F6449) as seen at right 
angles to its length (a). Points on the perimeter of an 
ellipse whose major and minor axes are the same 
lengths as the maximum and minimum widths of the 
cast (+). Ellipse whose major axis is the same length 
as that of the cast but twice its area in section (b). 
Circle with same perimeter as that of ellipse b (c). 

charge; Dr Alex Cook for his critical appraisal 
and support of the project; Natalie Caniilleri for 
her conversion of my rough drawings and 
diagrams into such fme Figures. 

LITERATURE CITED 

ANDERSON, J.M. & ANDERSON, H.M. 1985. 
Palaeoflora of Southern Africa- Prodromus of 
Southern African megafloras Devonian to Lower 
Cretaceous. (A. A. Balkema: Rotterdam). 

CAI, C. & WANQ Y. 1995. Pp. 1-689. In Li, X. (ed.) 
Fossil floras of China through the geological ages. 
(Guangdong Science & Technology Press: 
Quangzhou). 

CLIFFORD, H.T. 1996. Geometrical study of a cast of 
Leptophloeum ausn-ale (McCoy) Walton from 
Queensland. Memoirs of the Queensland 
Museum 39(2): 227-231. 

DAWSON, J.W. 1862. On the flora of the Devonian 
Period in Northeastern America. Quarterly 
Journal of the Geological Society, London: 
296-330. 

EDWARDS, D. & BERRY, C. 1991. Silurian and 
Devonian. Pp. 117-53. In Cleal, C.J. (ed.) Plant 
fossils in geological investigation. (Ellis 
Horwood: London). 

McCOY, F. 1874. Lepidodendron (Bergeria) australe 
(McCoy). Pp. 37-39. In Prodromus of the 
palaeontology of Victoria. Decade ! . (Geological 
Survey of Victoria: Melbourne). 

MORRIS, N. 1977. Carboniferous Jloras of eastern 
Australia, linpubl. PhD thesis, Universit>' of 
Newcastle, Australia. 

SCOTESE, C.R. & McKERROW, W.S. 1990. Revised 
world maps and Introduction. In McKerrow, W.S. 
& Sotese, C.R. (eds) Palaeozoic palacogeography 
and biogeography. Geological Society Memoir 
12: 1-21. 



APPENDIX 1 

Method of calculation using symbolic 
manipulator, Maple V Release 4. Maple V 
Release 3, 4 or 5 can be used for the deter- 
mination of the various calculations required in 
this paper and these problems are now discussed 
in general. Given the circumference of a circle, so 
that its diameter can be calculated, it is required to 
find the major and minor axes of ellipses which 
result from keeping their perimeters fixed as that 
of the original circle, but whose minor axes are 
reduced to some fraction of its radius. To find the 
perimeter of an ellipse it is necessary to calculate 
an elliptical integral which must be evaluated 
numerically. In fact, the perimeter is given by: 

P = 4 I " Va"cos ~u + b~sin "^u du 

where a and b are the semi-major and semi-minor 
axes of the ellipse. Maple allows the setting up of 
this integral in terms of a and b and, once the 
circumference of the original circle is ascer- 
tained, it is possible to calculate the semi-major 
axes of the ellipses whose semi-minor axes are 
reduced in a given ratio of their original value 
(Fig. 3). To do this a numerical method must be 
set up which starts with an initial approximation 
to the solution and then uses a steepest descent 
method to generate closer and closer approx- 
imations to the required solution. The degree of 
accuracy can be chosen as desired. Once the new 
ellipses have been found the positions of the 
margins of the leat-cushion bases must be 
calculated. Again this must be done numerically. 
If it is assumed that the leaves are equally spaced 
around the perimeters of the ellipses then, since 
the perimeters have been calculated, the arc lengths 
corresponding to the widths of the leaf-cushion 
bases are known — it was assumed there were 
eight leaves per quadrant (see Fig. 4). A similar 
steepest descent method is now used to step 
round the perimeter of the ellipse, successively 
finding the co-ordinates which define the 
margins of each leaf. This method was also used 
to produce Tables 2 and 3. Once the co-ordinates 
of these leaf-margins have been found, those on 
the contracted ellipses with the same major axes 
but smaller minor axes are found by merely 
holding their X co-ordinates constant and 
decreasing their Y co-ordinates in the appropriate 
ratios of the required contraction (see Fig. 4). 
Information on the Maple program can be 
obtained from M.R. Pemberton at e-mail mrp@ 
maths.uq.edu.au 



234 



MEMOIRS OF THE QUEENSLAND MUSEUM 



FOSSIL TYPHA IN AUSTRALIA. Memuirs of the 
Qucens/amJ Museum 45(2) ■ 234. 2000;- Recent invesligation 
ofoil shales recovered from a bore put down in the Casuarina 
Basin, about 25km SE of Rockhampton, Queensland has 
revealed seeds and fruits in the sediments. These were 
encountered in the course of a palynological study and would 
have otherwise been overlooked on account of their small 
size. In a forthcoming paper the seeds have been assigned to 
Typhaceae and the fmils to Restionaceae (Dettmann & 
Clifford, in press). Because seeds of Typhaceae are 
operculate it is important they not be confiised with moss 
capsules. The two are usually readily distinguished: unlike a 
seed the moss capsule is usually attached to a seta and has a 
peristome which is revealed when the operculum separates 
from the theca. However, il'ihe moss capsule has become 
detached from its seta and lacks a peristome it will resemble a 
Typha seed whose chalazal region has been damaged. These 
considerations led us to reconsider the identity of Muscites 
yullownesis Clifford & Cookson which was described on the 
basis of a single specimen isolated from a sample of brown 
coal (Clifford & Cookson. 1953) of Miocene age from Yal- 
loum (Blackburn & Sluiter. 1994). 

A comparison of the cell structure of the operculum of A/. 
yallournensis with that of the extant Typha domin^cnsis Pers. 
revealed no significant dilTerences (Fig. I ). Further support 
for the view thai the specimen of A/, yaiiournemis is a seed of 
Typha rather than a moss capsule is provided by the collar of 
cells from within which the operculum is shed and the ragged 
skirt of cells around the base of the supposed 'capsule'. Both 
of these are features ofTypha seeds and are clearly visible on 
the photograph of the holotv pe of A/, yallournensis. Accord- 
ingly, the species is here formally transferred lo that genus. 

Systematic Palaeobutiiny 

TYPHACEAh 
Tvpha yallournensis (Clifford & Cookson) comb. nov. 
(Fig. lA-C) 

Muscites yallournensis Clifford & Cookson, 1953: 54-55. 

MATERIAL. HOLOTYPE: NMV PI 5724; Latrobe Valley 
Coal Measures, Yallourn Scam; Miocene. 

REMARKS. There are no previous reports of Typha (type 
species T. latifolia L.) from the Australian fossil flora though 
MacPhail et al. ( 1994) and Blackburn & Sluiter ( 1994) report 
tlie presence of macro- and/or microfossils with affinities to 
Typhaceae and Sparganiaceae but make no positive ident- 
ifications to either. Therefore Typha yallournensis (Clifford 
& Cookson) comb. nov. becomes the first definite fossil 
record of the genus from Australia. 

Elsewhere fossil Typha seeds have been described from 
Maastrichian and vounger sediments of Europe (Chandler, 
1963; Collinson. 1983; licrendeen & Crane, 1995). Seeds of 
T. lalissima A. Braun closely resemble those of T. 
yallournensis but until the anatonn oiThe latter is known the 
two species cannot be regarded as conspecific. 

Acknowledgements 

We thank the Director of the Museum of Victoria and Dr S. 
McEouglilin. University of Melbourne for arranging access to 
the negative of the holot> pe ol" Typha yallournensis. 

Literature Cited 

t5l,ACKBURN, D.J. & SLUITER.. I.R.K. 1994, The Oligo-Miocene 
floras of south-eastern Australia. Pp. 328-367. In Hill. R.S. 
(ed.) Histor>' of the Australian vegetation: Cretaceous to 
Recent. (Cambridge Uni\ersit\ Press: Cambridge). 

CHANDLER, M.l:..l. 1963. Revision of the Oligocene tloras of tlie 
Isle of Wight. Bulletin of the British Museum (Natural History) 
Geology, London 6: 321-3S4. 

CLIFFORD, H.T. & COOKSON, l.C. 1953. Muscites yallournensis. a 




FIG. 1. Typha yallournensis (Clifford & Cookson) comb. 

nov. and T. domin^ensis Pers. A-C, T. yallournensis, 
holotype (NMV PI 5724). A, seed body showing chalazal 
region at base ( x 60). B-C. detached operculum { x 60 and 
X 120). D-(l, T. domin^ensis. D, E, Ci, QM F50036 seed 
showing partially detached operculum and chalazal region 
at base (D. x33. E. x 133, G, x 50). F, specimen QM 
F50037, detached operculum in lateral view ( x 133). 

fossil moss capsuk- from Yallourn, Vicloria. The Brvologlst 56; 
53-55. 

COLLINSON, M.E. 1983. Palaeoflorislic assemblages and 
paiaeoecology of the Lower Oligocene Bembridge Marls, 
Hamstead Ledge, Isle of Wight. Botanical Journal of the 
Linnean Society 86: 177-225. 

DHTTMANN, M.L, & CLIFFORD, H.T. in press. Monocotyledon fruits 
and seeds and an associated patynotlora from Eocene/ 
Oligocene sediments of coastal central Queensland. Australia. 
Review of Palaeobotany and Palynology. 

HERENDEEN. P.S. & CRANE, P.R. 1995. The fossil history of the 
monocotyledons. Pp. 1-21. In Rudall, P.J.. Cribb. P.J., Cribb, 
D.F- & Humphries, C..I. (eds) Monocotyledons: systematics 
and evolution. (Royal Botanic Gardens: Kew). 

MACPHAIL, M.K., ALLEY. MT., TRUSWELL, E.M. & SLUITER, 
LR.K. 1994. Early Tertiary' vegetation: evidence Irom spores 
and pollen. Pp.189-261. In Hill. R.S. (cd.) History of the 
Australian vegetation: Cretaceous to Recent. (Cambridge 
Lni\ersity Press; Cambridge). 

//. Trevor Clifford, Queensland Museum, PO Box 3300, 
Souih Brisbane 4101, Australia. Mary E. Dettmann. The 
University of Queensland, St Lucia, 4072, Australia. 11 
January 2000- 



TIL\CJEFriiSS3MM0MT!ir IJPPrR CARi^ONlFERQUS JERICHO FORMAtlON. 



;Al£XG COOK ANl^KBilRIfeBAW* 

GooK A.G. Bmi. K. 2000 0^ 30; Trace fbssiI^ fV.Hii the (.'ppc I .iibonifcraus JcricJio 
Formaliou, cenUaJ QueemUand. Memoin of the Queai&land Museum 45(2); 235-251. 
Brisbane. ISSN 0079-8835. 

A new iracf lu^^sll as^eililMa^ frorn lacustrine deposits of the Upper Carboniferous Jericho 
f otmatiocu ioe Jpe Gipup, Galilee Basin, Queensland^ is deseribed; it is similar to tiie 
Ifm&lozia ictulicdetissls found in other pan^ of Gondwanaland during the Laie^ 
Carbbtilierbusliq PprzQlsai^Jheiisseinbfage comprises e^cJusive^'oPieddlnKijfaraliel 
trades* interpreted as ^arttiropod locomotor)' traces, iTiainl> J^m&tiddf^^l^^^^^i |fihno5{>. 
nov., and Alphaichmts luphoensh ichnogen. ci ichnosp. nov. Other new taxa ate 
IsopotJichwts qiteemiantk'mk ichcnosp. nov., Bt^sophvct^s devh't ichnosp. nov. and 
Wadeichntis ntaryna i(^o£e]Uet i<4u(l(D9p* nov. □ Trace^jtHs, Carhonffemm wthtopod 
Iroifs, Qiiccm land- 
Alex G. Cook ti Kerrie Bonn, Queemiatid Mu^yeum. PO Hox .-^JOO. Saiiih Bmbatie 4 1 01, 



t)elicaifi ssni divetse trace fossil assemblages 
are ktiowtt from Laie Carboniferous to Eqriy 
P(^ian of Gondwanaland most notehly tlie DwyVa 
SerieSs South Africa (Savage. Anderson. 
1 ^75, 1576. 1 98 ] ), Soutli Amcncn ( R ocha-C. ainpos, 
1 967; Ac(ffft>laza& Buatois. 1 MVl , l Buatois 
& Mangano. 199.^, 1994. 19^)5) and Australia 
(Chapman, 1929; Gluct>sncT, 1957). Uniformity in 
these Gondwanan ichnofaunas allows recognition 
of jitter aU^ U^nfolozia, Isopadicbaus ^xki 
iMttTUk folinoGoenosfeS^ (Acteftolaiia & KiatoJs, 
1*>93) vvhicli are iisefu] biogeographic and broad 
biostratigraphic indicators, as well as rellecting 
widespread glacial conditions ortheir onset. This 
papet Is cpncemed with ^ell-pteserved trace 
fossils fhrmi sltialfctw quarries ot the Jtriello 
Pormatit^n alonu the Seduclbrd-AIpha Rd. SSF 
of Alpha, cLMiirul Queensland (Queensland 
Museum Locality QML993). 

The Jericho Formation d dominanll}/ 
fine-grained, siliciclastic unit forming part of the 
Joe Joe Group of tbe Galilee Basin. The unit was 
formal K' defined by Chay ^ Swarbnck (1975) 
who noted arthropod trails and suggested a 
I^UStrint? deposilional environment in associat- 
ion with periglacial ctmdiiions. Biosiraiigraphic 
control on the Joe Joe (iroiip was reviewed hy 
Jones Sl Truswcll ( UJ'^JZ) who plaeed llie Jericlio 
Formation in tbe latest Carboniferous ^.Steph- 
ijnianK poMdafftig the omet of glaciation. In 

order to elucidate the dcposiliunal conlexl for 
lllL•^c trace fossils wc examined jeco\crcd core 
iromlhe same seqacncc. 



DBPOSlTlONALSETTrNG 

txaminaiion of interval 770-740in in GSQ 
Jericho No. ! (see Swarbnck. 1974: fig. 2) 
revealed a lining upw ard sequence, interpreted as 
a deepening alluvial to lacustrine environment 
(Fig. 1). At the base of the unit, unbioturbated, 
medium- to coarse-grained pebbly sandstone 
represents alluvial deposition. Above, the 
sequence grades to iine-grained sandstone w iiha 
Jmcanmichnus iehnofabric and possible esc^i^: 
tr^s>. whioh we intprprel as a deltaic enviixnJ- 
ment Tlie secfuence grades progressively into a* 
biotMrbalcd, interbcdded sandstone and silistone 
unit with sinacresi^ cracks. c(^nt(>rlcd bedding 
and llaine siruclurcs, possibly relleeltng inllux of 
sedmient due to storms or seasonal input in a 
laeustrine environment. The upper portion of tlji^ 
uilit is ^oinposed of pinstriped siltstonem^ ve;^ 
fine-grained sandstone with Tamanadfa. This 
pari of Uie sequence v\e interpret to represent 
seasonal varvcs. The trace lo.ssii assemblage is 
interpreted as a Rusophycus ichnv)laeies. T he top 
7,5m of the section coniaini^ btoiurbaie4. imer- 
ho^ded^ecy fine-gcaiiied sandstone and silisttyne 
:md reflecrs deposition in a distal, t|uiet lacustrine 
env Lromneni. The sequence is overlain by a series 
pf Tjrograding deltaiadcpojsits* 

fetJSG&HVCtiS in this study 

tlie eponymous ichnogenera are i^t iialicified 
(following Bromley, 1996). because it %a fScics 

aiuinol an ichnotaxon that is tinder disctission {this 
is in line with the usage of la\a m biosnaligraphical 
zones^e^. Bi&pns zone). 



236 



MEMOIRS OF THE QUEENSLAND MUSEUM 



distal 
lacustrine 



proximal 
lacustrine 




El 



deltaic 



alluvial 



Key 




ball and pillow 
■* plant 

»>Tasmanidia-\ike 
tracks 



is 



synaerests 
cracks 



FIG. 1. Graphic log of GSQ Jericho No. 1 interval 
770-740m, showing relative depostional energy, 
interpreted depositional environments and ichno- 
fabrics adapted from Taylor & Goldring (1993). 

This assemblage falls into the Rusophycus 
ichnofacies that was suggested tentatively by 
Bromley (1996: 249) to represent a freshwater 
(Huvial and shallow lacustine) ichnofacies 
dominated by Repichnia and Cubichnia. 

In the Jericho Formation the Rusophycus 
ichnofacies is characterised by Repichnia, 
Cubichnia and Praedichnia (Table 1). 

Acenolaza & Buatois (1993) reviewed late 
Palaeozoic trace fossil assemblages of Argentina 
and identified four ichnocoenoses pertaining to 
differing lacustrine and fluvial sedimentary 
facies. They regarded the Umfolozia ichnocoen- 
osis as representing arthropod activity within a 
shallow lake, and the Isopodichnus ichnocoenosis 
as representing activity within ephemeral, shallow- 
pools, and channel and floodplain deposits. The 
Queensland material belongs within an Umfolozia 
ichnocoenosis, with arthropod crawling traces 
dominant. Keighley & Pickerill (1996) reviewed 
the interwoven taxonomic status of Isopodichus, 
Rusophycus and Cruziana. and provided a con- 
vincing argument against retaining Isopodichnus, 
placing ribbon like members of that taxon in 



Cruziana, and bilobate colTee-bean shaped 
members within Rusophycus. However Trewin 
( 1 976), Pollard ( 1 985) and Acenolaza & Buatois 
( 1 993) pointed out that Isopodichnus is generally 
smaller, with flaring ends. Furthermore the genus 
has a significant ecological and stratigraphic 
utility, and is here retained. 

FUNCTIONAL INTERPRETATION. Tlie absence 
of body fossils associated with the assemblage 
makes identification of the track-maker(s) 
difficult. The biserial nature of many of the traces 
indicates that they were made by arthropods. In 
Wadeichnus, interpreted antennae marks 
strengthens the arthropod identification. Groove 
markings on Isopodichnus queenslandensis, 
interpreted to have been made by telson drag, and 
the presence of paired scratch marks suggestive 
of at least 5 pairs of appendages on the related 
Rusophycus devisi, throws some light on the 
organisms responsible. The bifurcate distal 
appendage marks in Alphaichnus indicate that 
appendages were in the equally biramal 
(primitive) state, possibly representing activity 
of syncarid Crustacea. 

Tenninology follows Osgood ( 1 970) for 'trail', 
'track', 'pair' and imprint'. 'DistaP and 'prox- 
imal' refer to the distance from the axis of the 
trace fossil, 'medial' pertains to features within 
the axial zones of tracks and 'lateral' to those in 
the marginal zones. Most material is preserved in 
hyporelief and all descriptions should be read as 
such. No attempt is made to establish higher 
order systematics. 

SYSTEMATIC PALAEOICFINOLOGY 

Isopodichnus Bornemann, 1889 

TYPE SPECIES. Isopodichnus problematicus 
Bornemann 1889, from the Triassic of Germany. 

REMARKS. Isopodichnus is a problematic 
ichnogenus, and has been variably placed in 
Rusophycus and Cruziana, with considerable 
taxonomic confiision, further obfuscated by well 
known intergradation between the three 
ichnogenera (also see below). Full accounts and 
differing opinions on its resolution are given by 
Osgood (1970), Hantzschel (1975) Keighley & 
Pickerill (1996) and Bromley (1996). The 
problems are intertwined and we do not propose 
to resolve this significant controversy. Bromley 
( 1 996: 1 84) provided the most accurate summary 
of this nomenclatural debate: 'I carmot see how 
Isopodichnus can be considered available other 
than a muddled junior synonym of both Cruzicma 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



237 



TABLE \. Ethological classification of ichnolaxa 
described from the Jericho Fmn near Alpha. 



Ethological 
classification 


Ichnotaxon 


Figure numbers 


Repichnia 


hopodichnus 
queenslcmdicus 


Figs 2A,B. 13B 


Cubichnia ! Rmophycus devisi 


Fig- 3 


Repichnia 


Tasmanadia 
glaessneri 


Figs 6. 7 


Repichnia with 
Praedichnia 


Wadeichnus maryae 


Fig. 4 


Repichnia with 
Praedichnia 


Alphaichrtus 
alphaensis 


Figs 8, 9. I OA, 12. 
?10B,C 


?Fugichnia 


Indet. sweep and 
scurry marks 


Fig. 13D 


?Praedichnia 


Indel- paired ap- 
pendage marks 


Fig. 12 



and Rusophycus\ The taxonomic stand of 
Keighley & Pickerill (1996) would have 
Isopodichnm queemlandensis within Cruzianu. 
We retain Isopodichmis here as a genus-of- 
convenience awaiting resolution of the tripartite 
nomenclatural confiision. We note the consider- 
able difference in the classic Early Palaeozoic 
Cruziana morphology and this material. The 
dichotomy between type specimens of Cruziana 
and those of Isopodichnus must be further 
investigated. 

Isopodichnus queenslandensis ichnosp. nov. 
(Figs 2A,B, 11 (part), 12B) 

ETYMOLOGY. From Queensland. 

MATERIAL. HOLOTYPE: QMF32233. PARAT\TES: 
QMF34026, 39062 all from QML993. 

DIAGNOSIS. Exclusively elongate and contin- 
uous Isopodichnus. 

DESCRIPTION. Straight, gently curved or 
weakly sinuous ribbon-like trail, up to 7mm wide 
consisting of (in hyporeliel) 2 parallel marginal 
longitudinal fiirrows and a central (axial) channel 
containing up to 6 subparallel longitudinal striae. 
Axial channel occupying slightly over Vi track 
width, with numerous fine longittidinal threads 
and a weak central ridge deviating from the 
mid-line along the length of the trail. Outermost 
longitudinal ridge in the axis with sporadic but 
numerous oblique, short striae. Marginal ridges 
with fine obliquely transverse striae, extending 
sub-perpendicularly beyond the edge of the trail. 
In some specimens these striae bifurcate, 
particularly as the trail shallows. There are 
numerous tiny (I -2mm wide) examples of the 
track (Fig. 9, large arrow) preser\^ed in epirelief, 



the marginal parts of the track are slightly more 
pronounced than in larger specimens. 

REMARKS. The central threads represent the 
drag marks of a telson with furcae, with the 
sporadic striae representing setae on the furcae. 
Short imprints on the edge of the track represent 
appendage marks. The relative depth of the track 
reflects the softness of the muddy substrate. Tlie 
elongate nature of the trace and the internal striae 
separate this ichnospecies from material figured 
Isopodichnus oshornei GlaQssncr^ 1957: pi. U), 
fig. 2a,b (partim); fig. 3, pi. 1 1 figs 1-3) from the 
Carboniferous near Seaham, NSW. The holotype 
of Isopodichnus queenslandensis is associated 
with Rusophycus. Smaller representatives of this 
taxon are interpreted as representing juveniles. 

Rusophycus Hall, 1852 

TYPE SPECIES. Significant problems with the nominate 

type ichnospecies are still lo be resolved (Keighlev & 
Pickerill, 1996). 

REMARKS. Osgood (1970) restricted Isopod- 
ichnus Bomemann to small Rusophycus-Wkc 
imprints of non-trilobite origin as well as to those 
of trilobite origin. Hantzschel (1975) suggested 
that Rusophycus be restricted to identifiable 
trilobite resting traces based on a genetic and 
stratigraphic methodology rather than an 
ethological/morphological approach. Use of Iso- 
podichnus for small short traces w as discussed bv 
Glaessner (1957), Osaood (1970), Birkenmajer 
& Bruton ( 1 97 1) and Hantzschel ( 1 975). Keighley 
& Pickerill (1996) argued against Isopodichnus 
and placed bilohate coffee-bean shaped members 
in Rusophycus. Buatois & Mangano (1993) 
followed this using Rusophycus for non-marine, 
late Palaeozoic traces with this short bilobate 
morphology. 

In the Alpha material Isopodichnus queens- 
landensis and Rusophycus devisi are associated 
and intergrade (Fig. 3C). 

Rusophycus devisi ichnosp. nov. 

(Tigs 3, 7A(part)) 

ETYMOLOGY For C.W. De Yis, who described the first 
trace fossils from Queensland (Oe Vis, 191 1). 

MATERIAL. HOLOTYPE: QMF32232. PARATYPES: 
QMF34026 (several specimens). 34090, 34069 (several 
specimens), all from QML993. 

DIAGNOSIS. Small, elongate longitudinally 
subsymmetrical to irregular bilobate trace, 
show ing at least 5 cun ed paired ridges in hypo- 
relief within each lobe. 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



239 



■€ ) 



V* 



B 




4 



D 



FIG. 3. Riisophycus devisi ichnogen. et ichnosp. nov.; A, QMF32232 holotype X 1 ; B, F34026 X 1; C, F34026, 
another track on of same slab; D, F32233, Riisophyciis devisi ichnogen. et ichnosp. nov. in association with 
Isopodichniis queenslandensis ichnosp. nov. X 1 . 



DESCRIPTION. Small trace up to 11mm wide 
and 13mm long, 2 lobes subsymmetrical about a 
midline or irregular. Each lobe with at least 5 
arcuate transverse ridges of which some are 
slightly divergent. Midzone of trace with an 
arcuate depression in hyporelief 



REMARKS. QMF32232 has 5 parts to this trace, 
2 of which are coalesced (Fig. 3A) representing 
saltation-style locomotion between short resting 
traces. QMF34026 has 2 small asymmetrical 
specimens and an undertracked specimen of a 
transitional trace between Rusophycus and 



240 



MhMOIRS OF THE QUEENSLAND MUSEUM 




TRACE FOSSILS FROM CENTRAL QUEENSLAND 



241 



Isopodichnus (Fig. 3B). QMF32233 shows Ruso- 
phyciis in association with Isopodichmts (Fig. 
3D) where the track may have been utilised twice 
by the same type of trace-producing organism. 
Other specimens are associated with Tasmanadia. 

Tasmanadia Chapman, 1929 

TYPE SPECIES. Tasmanadia twelvetreeemis Chapman, 
1929 from the Upper Carboniferous Wynyard Tillite, 
Tasmania. 

REMARKS. Chapman (1929) erected Tasman- 
adia for purported Cambrian annelid body fossils, 
reinterpreted to be Carboniferous arthropod 
tracks by Glaessner (1957) and Gulline (1967). 
Bromley & Asgaard (1979) regarded Tasman- 
adia as a junior synonym of Dipliclmites but 
contrarily indicated significant differences in the 
fine morphology of the tracks; their extreme 
Mumping' view where many arthropod track genera 
were synonymised is not here adopted. 

Tasmanadia glaessneri ichnosp. nov. 
(Figs 6, 7A (part)) 

ETYMOLOGY. For the late M. R Glaessner. 

MATERIAL. HOLOTYPE: QMF39069. PARATYPES: 
QMF32229, 34054, 34060, 34065, 34081, 34083, 34088, 
34090, all from QML993. 

DIAGNOSIS. Biserial track 8-1 1mm wide, of 2 
near-symmetrical rows of fine ridges, nearly 
perpendicular to the track axis, in close, slightly 
divergent pairs. 

DESCRIPTION. Track biserial, elongate, gently 
curved, of near symmetrical rows of fme ridges in 
hyporelief, 8-1 1mm in total width, with 3-4mm 
between inner ends of rows of ridges. Fine ridges 
in closely spaced pairs diverging slightly or 
subparallel, individually very weakly arcuate, 
perpendicular to the track axis. Sporadically 
finer, more strongly divergent appendage marks 
marginal to the track pairs. Rare lengthwise, but 
short marks along the track axis. Some specimens 
over 500mm long. 

REMARKS. The paired track marks indicate 
Tasmanadia. This material is differentiated fi^om 
the type species by the greater divergence of the 
distal ends of the ridges forming the track pairs. 
Permichniuw Guthorl, 1934 has slightly more 
divergent paired ridges, but they emerge fi"om 
subcircular foot impressions, lacking in this 
material. Maculichna Anderson, 1975 possesses 
rows of paired spots, and lacks the paired 
divergent elongate marks of this material. 




I 



antenna mark 




/ pleopod withdrawal 




FIG. 5. Interpreted schema for Wadeichmis maryae 
ichnogen. el ichnosp. nov. based on holotype 
specimen, x2. 

Umfolozia Savage 1971 also possesses a series of 
paired and complexly arranged dots in contrast to 
the simplicity of this material. Specimens of this 
ichnotaxa grade into Rusophycus and Alphaich- 
mis, the latter evidenced by the rare occurrence of 
fine marks adjacent to the paired ridges. Shallow 
undertracking suggests greater substrate 
firmness than for Alphaichnus md Isopodichnus. 

Alphaichnus ichnogen. nov. 

TYPE SPECIES. Alphaichnus alphaensis ichnogen. et sp. 
nov. 

DIAGNOSIS. Biserial, gently curved, to straight 
trace consisting of subsymmetrica! rows of up to 
three sets of paired ridges which are distally 



242 



MEMOIRS Oi' THE QUEENSLAND MUSEUM 




Fia 6. Tasmanadia glacssmn ichnosp. nov., spocii]jens f^om slab QMF3409Q. A, Qvefprintei liyp<?i;elief 
specjinens^aiadeteitoinafcivveep^^ x l-B, further detail specimen xlT 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



243 




FIG. 7. A, Tasmanadia glaessneri inchosp. nov., QMF39069 holotype, (arrow) with associated Rusophycus devisi 
ichnosp. nov. and indeterminate traces x L B, indet. appendage marks, QMF34067 x 2. 



244 



MEMOIRS Of THE QUEENSLAND MUSEUM 




FIG 8. Alphaichms alphaemis ichnogen. el ichnosp. nov. QMF34068 x 2. B, holotype F34084 x 2; C,F34084, 
track on different part of same slab x 2. 



TRACE FOSSILS FROM CENTR.\L QUEENSLAND 



245 



branched and have short fine striae. Axis with or 
without arcuate ridges or transverse oblique ridges. 

Alphaichnus alphaensis ichnogen. et sp. nov. 
(Figs 8-11) 

ETYMOLOGY. From the town of Alpha 

MATERIAL. HOLOTYPE: QMF34084. PARATYPES: 

QMF32231, 34033, 34039, 34043, 34057, 34059, 
34066-34068, 34072. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Long, straight to gently curved, 
irregular biserial trace, without closing loops, up 
to 1 8mm outer width, 8mm inner width. Internal 
organisation complex of subparallel track sets of 
up to 3 pairs of variable length transverse ridges, 
each of which may be distally bifurcate or 
trifUrcate, with short, very fine longitudinal striae 
perpendicular to their length. Length of 
individual ridges in each set increasing towards 
the margin of the track. Central zone of the track 
with up to 3 distinct or 2 axially connected, 
curved transverse ridges, or a series of up to 4 
irregularly disposed oblique short ridges. On 
some specimens additional asymmetrical curved 
transverse ridges at the margins of the track. 
Individual pairs highly variable, but approach 
symmetry when the trace is nearly straight. 
Longitudinal striae, oblique and arcuate ridges 
sporadic along length of track, within the axial 
zone. In some specimens medial arcuate, sym- 
metrical ridges, with tiny longitudinal striae with 
a series of adaxially disposed divergent ridges at 
their margins, and a number of lateral short marks 
fanning around the individual paired ridges. 

REMARKS. There are a large number of biserial 
arthropod tracks superficially similar to this ma- 
terial, but most lack the internal variablity and 
complexity of this ichnotaxon, particularly 
within the track axis. Diplichnites govenderi Sav- 
age, 1971, lacks the flaring and branching of 
individual tracks, lacks medial ridges and is, in 
general, a simpler trace. The considerable varia- 
tion in this genus reflects similar morphological 
variation encountered by Anderson (1975) in 
Umfolozia Savage, 1971, but Umfolozia bears 
appendage marks and sinuously arranged oval 
marks in the medial zone and lacks the complex- 
ity of medial arcs and branching ridges present in 
Alphaichnus. 

Walter (1983) discussed many late Palaeozoic 
ichnotaxa attributed to arthropods. Heterotripo- 
dichtms divaricatus Walter and H. longitarsalis 
Walter approach the present material in 



complexity, but in both, the appendage marks are 
more longitudinally arranged within the trace. 

Irregular morphology makes the identification 
of repeat distances difficult. Repetition of medial 
ridges is interspaced by 5 track pairs, which 
suggests minimum track repetition. Track pairs 
are interpreted as appendage marks with their 
distal biftircalions and other branchings repres- 
enting the 2 roughly equally-sized rami. Medial 
oval and arcuate marks are interpreted as pleopod 
imprints and withdrawal marks. Longitudinal striae 
on the appendage marks setae. Distal fine arcuate 
marks are interpreted as traces of antennae. 

The differing depth of track penetration 
displayed by the many slabs examined with 
undertracks suggests that substrate fiminess was 
variable, but was high in the case of generation of 
this track compared to the occurrences of Iso- 
podichms queenslandensis. 

Wadeichnus ichnogen. Nov. 

TYPE SPECIES. Wadeichnus maryae ichnogen et 
ichnosp. nov. 

ETYMOLOGY. For Mary Wade. 

DIAGNOSIS. Biserial track of elongate, lobate, 
longitudinally disposed marks, consisting of up 
to 5 elongate striae, terminating in a horseshoe- 
shaped depression within which the striae fan 
slightly; with or without marginal arcuate ridges 
perpendicular to track axis, and with or without a 
bounding pair of longitudinal fine ridges. 

Wadeichnus maryae ichnogen. et ichnosp. nov. 
(Figs2C,D, 4,5, 12C) 

MATERIAL. HOLOTYPE: QMF39063. PARATYPES: 
QMF39065, 3906L 

ETYMOLOGY. For Mao Wade. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Biserial, somewhat discontin- 
uous track, up to 14mm wide, of symmetrical or 
subsymmetrical lobate marks, up to 8mm long, of 
3-5 longitudinal striae which diverge slightly 
within a horseshoe-shaped depression, 
accompanied by marginal fine ridges, papillate, 
which arc perpendicular to the trace axis and a 
pair of longitudinal fine threads at the margin of 
the trace. Some specimens show a central thread 
running lengthwise along the trace. In two 
specimens the orientation of the appendage 
marks (striae sets) are oblique to the trace axis. 



246 



MEMOIRS OF THE QUEENSLAND MUSEUM 



B 



FIG. 9. A, Alphaichnus alphaensis ichnogen. et ichnosp. nov. QMF3223 1 x 1 . B, C, cf. Alphaicfmus. QMF34036 
and F39064 respectively. 



REMARKS. The arcuate ridges perpendicular to 
the axis are interpreted as antennae marks 
elongate striation pairs as pleopod marks and 
horseshoe depressions (in hyporehef) interpreted 
as withdrawal markings (sensu Osgood, 1970). 
The sporadically present central thread may be a 
telson drag mark. It is likely that the specimen 
assigned to cf. Alphaichnus (Fig. 9B,C) is 



transitional between Alphaichnus and this 
ichnotaxon. Wadelchnus probably was formed on 
a semifirm substrate with only minimal contact 
of the arthropod body and the surface sediment. 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



247 




FIG. 10. Alphaichnus alphaensis ichnogen. et ichnosp. nov. QMF39068 X L 



indet. appendage marks 

(Fig. 7B) 

MATERIAL. QMF34067. 

DESCRIPTION. Small elongate series, 11mm 
long and 5mm wide of arcuate or lunulate to 
V-shaped wrinkle marks in hyporelief, up to 6 in 



an individual series, with discontinuous sets in 
close association. 

REMARKS. These marks probably represent 
asymmetrical appendage falls upon the substrate 
with subsequent, but penecontemporaneous 
plastic deformation, probably on a low slope. The 
lack of material available and the unusual 



248 



MEMOIRS OF THE QUEENSLAND MUSEUM 



■ >• "It* 



FIG. 1 1 . Alphaichnus alphaensis ichnogen. et ichnosp. nov. QMF34084, hololype (small arrow) part of composite 
slab, X 1. Isopodichmis queensiondensis, juvQmh specimens (large arrow). Note the numerous indeterminate 
appendage marks (open triangles). 



morphology prevents any accurate assignment. 
Anderson (1975) described bedding surface 
slump structures similar to those found in the 
Alpha material. 

indet. paired appendage marks 

(Fig. 11 (part)) 



DESCRIPTION. Almost all slabs containing 
Wadeichnus and Alphaichnus have isolated, 
shallow small paired or more rarely single holes 
(in epirelief). They are commonly triangular with 
one apex deeper, or they are subquadrate with no 
directional shallowing. Pairs are separated by 
2-3mm. No continuity in sets of pairs can be 
identified. 



MATERIAL. Part of slab with QMF34084. 



REMARKS. We interpret these holes as 
appendage pluck-out or withdrawal marks 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



249 




FIG. 1 2. A, Alphaichnus alphaensis ichnogen. et ichnosp. nov., and indeterminate 'sweep' mark, QMF34066 X 2. 
B, Isopodichms qiieemlandemis ichnosp. nov. QMF39062, showing sinuosity, x 1. C, Wadeichnus niaryae 
ichnogen. et ichnosp. nov. QMF3906I x2. D, indeterminate scurr>' marks QMF39067 X 1. 



250 



MEMOIRS OF THE QUEENSLAND MUSEUM 



associated with isolated substrate interactions. 
The association of these holes with the more 
complex traces shows differing behaviour 
between the maker of Alphaicbnus and the 
originator of these marks, but we cannot 
determine whether it would be a similar animal. 

indeterminate 'sweep' marks 
(Fig. 6 (part)) 

MATERIAL. OMF34090 

REMARKS. On a number of specimens are 
elongate curved and recurved ridges which lack 
repetitive association or are in groups of 1 or 2. 
These ridges are up to 25mm long, and <lmm 
thick, gently curved with a stepped weak bilateral 
symmetry, if at all. These structures are 
interpreted as sweep marks from some organism; 
they are much too small and non-persistant to be 
Umlichnia (sensu Anderson, 1976; Buatois & 
Mangano, 1994) and are indeterminate. The 
material resembles StiaJia pillosa Smith of 
Walker (1985), but lacks the abundant scratch 
marks. 

indeterminate scurry marks 

(Fig. 12D) 

MATERIAL. QMF 39067. 

REMARKS. Small, disordered set of weakly to 
strongly divergent ridges with an impersistent, 
marginal sinuous set of bordering threads. 
Imprints (ridges in hyporelief) are in groups of 3 
or more and are short, sporadically divided and 
bear fme transverse striae. The specimen is 
interpreted as a set of arthropod pleopod marks 
where the pleopods bear setae and are probably 
biramal. The trace architecture is inferred as 
resulting from an arthropod scurrying across the 
firm substrate. The sinuous threads are 
interpreted as antennae marks. 

ACKNOWLEDGEMENTS 

We thank Mary Wade and Peter Jell for their 
assistance and encouragment; Luis Buatois & 
Gabriela Mangano. Geological Survey of 
Kansas, Ron Pickerill, University of New 
Brunswick for providing important literature; 
Tony Wright for topotype material of 
Isopodichmis osbornei from Scaham; Natalie 
Camilleri, Paul Tiemey, Chris & Dawn Schnur, 
Terry Smith and Colin McHenry are thanked for 
assistance in the field; Jeff Wright is thanked for 
his patience with photographs. Amy Robinson 



for her assistance with preparation, Bany Webby 
for helpfril criticism on the manuscript. 

LITERATURE CITED 

ACENOLAZA, F.G & BUATOIS, L.A. 1991. Trazas 
fosiles de Palcozoico superioir continental 
argentino. Ameghiniana 28: 89-108. 

1993. Nonmarine perigondwanic trace fossils from 
the late Paleozoic of Argentina. Ichnos 2: 
183-201. 

ANDERSON, A.M. 1975. Turbidites and artliropod 
trackways in the Dwyka glacial deposits (early 
Pemiian) of southern Africa. Transactions of the 
Geological Society of South Africa 78(2): 
265-273 

1976. Fish trails from the Early Permian of South 

Africa. Palaeontology 19: 397-409. 
1981. The Umfolozia arthropod trackways in the 

Permian and Ecca Series of South Africa. .lournal 

of Paleontology 55: 265-273. 
BIRKEMAJER, \C. 8l BRUTON, D.L. 1971. Some 

trilobite resting and crawling traces. Lethaia 4: 

303-319. 

BORNEMANN, J.G 1889. Uber den Buntsandstein in 
Deutschland und seine Bedeutung fur die Trias. 
Beitrage, Geologic , Palaontologie 1: 1-61. 

BROMLEY, R.G. 1996. Trace Fossils: Biology, 
taphonomy and applications. (2nd edn). 
(Chapman & Hall: London). 

BROMLEY, R.G & ASGARD, U, 1979. Triassic 
IVeshwater ichnococnoses from Carlsberg Fjord, 
east Greenland. Palaeogeography, Palaeocliniat- 
ology, Palaeoecology 28: 39-80. 

BUATOIS, L.A. & MANGANO, M.G 1993. Trace 
fossils from a Carboniferous turbiditic lake: 
implications for the recognition of additional 
non-marine facics. Ichnos 2: 237-258 

1994. Pistas de peces en el carbonifero de la cuenca 
Paganzo (Argentina): su significado 
estratitirafico y paleoambiental. Ameghiniana 
31: 33-40. 

1995. The paleoenvironmenial and palaeo- 
ecological significance of the lacustrine Mcrmia 
ichnofacies: an archetypical subaqueous 
non-marine trace fossil assemblage. Ichnos 4: 
151-161. 

CHAPMAN, P. 1929. On some remarkable annelid 
remains from the Arthur River, N.W. Tasmania. 
Proceedings of the Royal Societ\' of Tasmania 
1928: 1-5. 

DE VIS, C.W. 1911. Annelid trails. Annals of the 

Queensland Museum 10: 12-14. 
GLAESSNER, M.F. 1957. Palaeozoic arthropod trails 

from Australia. Palaontologische Zeitschrift. 31: 

103-109. 

GRAY, A.R.G & SWARBRICK, C.F.J. 1975. Nomen- 
clature of late Palaeozoic strata in the northeastem 
Galilee Basin. Queensland Government Mining 
Joumai 76: 344-352. 



TRACE FOSSILS FROM CENTRAL QUEENSLAND 



251 



GULLINH. A.B. 1%7. The llrst proved Carboniferous 
dept)sils in lasmania. Australian Journal of 
Science 29: 332-333. 

GUTHORL, R 1 934. Die Arthropoden aus dein Carbon 
und Perm des Saar-Nahe-Pfalz-Gebeites. 
Abliandlungen der Preussischen Geolo^ischen 
Latidesaiistall 164: 1-219. 

HALL. J. 1852. Palaeontolgv of New York Vol. 2. 
(Albaii\; \e\v York). 

HANZSCHEL, W. 1962, Trace fossils and Probletn- 
attca. Pp. W177-245. In Moor^ R.C, (ed,) 
Treatise on invertebi^ paleontology. Part W. 
(Geological Society of At&erica & Umvasity of 
Kansas Press: New Yoife & Lawimce). 
1975. TVace Fossils and Problematica (2nd edn). 
Part W. Misccllania Supplement \. in Teicherl, 
C. (ed.) Treatise on Invertebrate Paleontology. 
( Geological Society of America & L'niversity of 
Kansas Press: Boulder. Colorado & Lawrence, 
Kansas I. 

JONES. M.J. & TRLISWLLL, E.M. 1992. Late 
Carboniterous and Faii> Permian paKnostrai- 
igrapln of the Joe .loe Group, southern Cialilee 
Basin, Queensland, and i m p I i e a t i <:) n s f o r 
Gondwanan slraligrapln. BMR .lourna!, Geologv 
and Geophvsics 13(2): 143-1S5. 

KEIGHLEY; D.A. <& PICRERILL. R.tC 1996. Small 
Cruziamu RusophycuSy and related ichnotaxa 
from eastern Canada: the nomenclatural debate 
and s\siematic ichnolog>', Ichnos4: 261-285 

OSGOOD, R.a 1970, Trace fbssite of *e Cincinnati 
^oea/Palaeontc^phica Americana. 6: 281-439. 



POLLARD, J.E. 1985. Isupodichmis. related arthropod 
trace fossils and notosiraceans tmm Triassic 
fluNial sediments. Iransactions of the Royal 
Society of Edinburgh, Earth Sciences 76: 
273-285, 

ROCHA-CHAMPOS, A.C. 1967. TheTubarao Group 
in the Brazilian portion of the Parana Basin. Pp. 

27-102. In Bigarella. J.J.. Becker. K.D. & Pinto. 
I.D. (eds) Problems in Brazilian (iondwana 
Geology. (Curiliba: Brazil). 

SAVAGE, N.M. 197 L A varvite ichnocoenosis from the 
Dwyka series ofNatal. Lethaia4: 217-233. 

SWARBRICK, C.F.J. 1974. Stratigraphic iliili iL 
report- QSQ Jericho 1. Queensland Govenimcm 
Minu^lournal 75(872): 210-215. 

TAYLOR, AJW. & GOLDRING, R. 1993. Desciiption 
and anals^is of biotiirbaii«>n and tchno^bdCf 
Jbum^ of the Geological Society, London 150: 
14!-148. 

TREWIN. N.ll. l^>7o. hopodichnus in a new nacc 
fossll assemblage Ironi the Old Red Sandstone. 
Lethaia 9:29-37. 

WAl.I KFR. R G. l'^.S4. Facics Models. (Gieoscience 

Canada: Foronto). 

WALTER, H. 1983. Zur Taxonomie, Okologie und 
Bioslratigraphie der Ichnia limnisch- 
terresirischer Arthropoden des mittcleuro- 
paischen Jungpalaozoikums. Freiberget 
Forschm^shefteReihe C 382: 146-193. 



252 



MEMOIRS OF THE QUEENSLAND MUSEUM 



AUSTRAIONEMA FROM THE SILURIAN OF THE 
YASS BASIN, NEW SOUTH WALES. Memoirs of the 
Queens/and Museum 45(2): 252. 2000:- The g>'ronematine 
gastropod Austrahnema Tassell has hitherto not been 
recorded from the Silurian of Austraha despite its abundance 
in AuslraHan Early Devonian gastropod faunas (Tassell, 
1980) and its presence in the Silurian elsewhere (Gubanov & 
Yochelson, 1994). Aiistralonemu recovered from the 
Barrandella Shales of the Yass Basin NSW extends the range 
of the genus within Australia. It confirms that Silurian 
Austrahnema belong to base stock of Gyronematinae, whose 
plesiomorphic state is characterised by one order of cords, 
lacking nodes. The specimen here described was recovered 
from the Barrandella Shales, Hattons Corner near Yass NSW. 
Recent work, summarised by Strusz (1989: 17), assigned a 
Ludlovian (Gorstian) age to the Barrandella Shale. This 
occurrence establishes the presence of basal stock of this 
subfamily in Australia prior to a significant Devonian 
diversification of gyronematines. 

Systematic Palaeontology 

ARCHAEOGASTROPODA 
TROCHIINA Cox & Knight, I960 
PLATYCERATOIDEA Mall, 1859 

HOLOPEIDAE Wenz, 1938 
GYRONEMATINAE Knight, 1956 

Australonema sp. 

(Fig.l) 

Material. QMF40825. Collected A.J. Wright., Barrandella 
Shales, Hattons Corner, Yass, NSW. 
Description. Large turbiniforin shell 41mm high, 33mm 
wide, anomphalous, sutures impressed. Shell profile well 
rounded, meeting suture at an angle of approximately 15° 
from horizontal. Mid-whorl at peripher>-. Upper whorl face 
bears 4 strong cords which lack nodes. Lower whorl face with 
at least 1 cord, but imperfectly preserved lower face ornament 
otherwise. Growth lines are fine and numerous, orthocline to 
ver>' slightly opisthocline. Protoconch unknown. 
Remarks. The specimen is noteworthy but I am reluctant to 
erect a new species based on a single occurrence. Aust- 
ralonema australis (Etheridge Jr, 1890) from the Lihdale 
Limestone has more numerous spiral cords. The specimen is 
closest to .4ustralonema sp. A of Tassell (1980) from the 
Silurian of Dudley, UK, but it also has a greater number of 
cords on the whorl face. 



Acknowledgements 

I thank Tony Wright for forwarding the specimen and 
Natalie Camilleri for photography. 

Literature Cited 

TASSELL, C.B. 1980. Further gastropods from the Early Devonian 

Lilydale Limestone, Victoria. Records of the Queen Victoria 

Museum 69: 3-27. 
ETHERIDGE Jr R. 1 890. Descriptions of Upper Silurian fossils from 

the Lilydale Limestone, upper Yarra district, Victoria. Records 

of the Australian Museum 1(3): 60-67. 
GLiBANOV. A.P. & YOCHELSON, E.L. 1994. A Wenlockian 

(Silurian) gastropod shell and operculum from Siberia. Journal 

of Palaeontology 68(3): 486-491. 
STRUSZ. D.L.I 989. Australian Phanerozoic Timescales 3: Silurian. 

Bureau of Mineral Resources, Geology and Geophysics 

Record 1989/33: 1-28. 

Alex G. Cook., Queensland Museum, PO Box 3300, South 
Brisbane 4101, Australia: 10 September 1999. 




FIG. 1. Australonema sp. QMF40825. xl; A, apertural 
view; B, side view. 



AUSTR.\LIAN LEAP-TAILED GECKOS: PHYLOGENY, ANEW GENUS, TWO NEW 

SPECIES AND OTHER NEW DATA 



PJ. COUPER, C.J. SCHNEIDER, CJ. HOSKIN AND J.A. COVACEVICH 

Coiiper, P.J., Schneider, CJ., Hoskin, C.J. & Covacevich, JA. 2000 06 30: Australian 
leaf-lailed geckos: phylogeny, a new genus, two new species and other new data. Memoirs of 
the Queemland Museum 45(2): 253-265. Brisbane. ISSN 0079-8835. 

Phylogenetic analyses of the leaf-tailed geckos, based on DNA sequences from the 
mitochondrial cytochrome b gene, show that Saltuarius, as currently constittited, is 
paraphyletic. Saltuarius comutus, S. salehrosus, S. wyioerba and S. swaini form a 
weil-supported monophyletic group which is the sister group to the apparently monophyletic 
Phyllurus. 'Saltuarius^ occultus is tlie sister group to the clade containing Phyllurus and all 
other Saltuarius spp. Thus, 'Saltuarius' occultus represents a long, independent, 
evolutionary lineage within the leaf-tailed geckos and is recognised from both 
morphological and molecular data as dislincl at the generic level (Orraya gen. nov.). Orraya 
gen. nov. can be distinguished from all other Australian padless carphodactylines by a 
combination of four apomorphies, the most obvious being greatly elongated cei"vical 
vertebrae. The phylogenetic analyses also revealed genetically distinct populations from 
Chaelundi State Forest, New^ South Wales and Oakview State Forest, southeast Queensland. 
These specimens are assigned to S. wyhcrha and R caudiarmulatus respectively, pending 
examination of more material. Description of Phyllurus amnicola sp. nov. and P. 
championae sp. nov. brings to 12 the number of leaf-tailed geckos from eastern Australia. 
The former, known only from Mt Elliot, NEQ, is separated from its congeners by a 
combination of characters (large size; very small, spinose body tubercules; a leaf-shaped tail 
with the anterior-mosl band broken, but spanning full tail width; and a partially-divided 
rostral with 5-6 scales along its dorsal margin). Phyllurus championae sp. nov. from only two 
localities (Cameron Ck and Blue Mtn, MEQ), is the only Phyllurus species with a 
leaf-shaped tail and a fully divided rostral scale. Large genetic distances between P amnicola 
sp. nov. , P. championae sp. nov. and previously recognised Phyllurus spp. further support the 
recognition of these species. They join a long list of rainforest reptile taxa known from only 
single localities or very narrow ranges. The distribution of Meaf-lails' reflects the 
preservation of ancient taxa in relictual rainforest and elevated heath fragments in eastern 
Australia. Despite these narrow ranges, and because all collection localities for leaf-tails are 
in reserves, for conservation purposes under lUCN definitions, they should be classed 'Data 
Detlcicni'. □ Reptilia. Gekkonidae, Phyllurus spp., Saltuarius spp., Orraya gen. nov.; 
phylogeny; eastern Australia. 

Patrick Couper Jeanefte Covacevich. Queensland Museum. PO Box 3300. South Brisbane 
4101. Australia; Christopher J. Schneider. Department of Biology, Boston University, 
Boston MA 02215, USA; Conrad Hoskin. Department of Zoology^ & Entomology, University 
of Queensland, St Lucia 4072. Australia; 7 February- 2000. 



Australia's rainforests and adjoining moist 
sclerophyll forests and heaths are well known for 
their high diversity and for many species con- 
fmed to either single localities, or very narrow 
ranges. Leaf-lailed geckos from such forests well 
illustrate these characteristics. For nearly 200 
years of discovery and description of Australia's 
reptiles, only two species of 'leaf-tails', 
Phyllurus platurus (Shaw, 1 790) and P. cormitus 
(Ogilby, 1892) = Saltuarius cornutus 
(Ogilby,1892), were known. Morphological 
studies since 1975 have resulted in the 
recognition of many new species and the genus 
Saltuarius Couper, Covacevich & Moritz, 1993 
— P. caudiarmulatus Covacevich, 1975; P. isis 



Couper, Covacevich & Moritz, 1993; P. nepthys 
Couper, Covacevich & Moritz, 1993; P. ossa 
Couper, Covacevich & Moritz, 1993; S. 
salehrosus (Covacevich. 1975); Saltuarius 
occultus Couper, Covacevich & Moritz, 1993 
and S. .swaini (Wells & Wellington, 1985). Eight 
of the 12 presently known species have narrow 
distributions with four confined lo single 
localities. Molecular analyses, which have both 
confirmed and refined previous morpho- 
logically-based taxonomic work on these padless 
Australian carphodactyline geckos, commenced 
in the mid-1990s. For the first time, a 
combination of both methods was used in the 
description of S. wyberba Couper, Schneider & 



254 



MEMOIRS OF THE QUEENSLAND MUSEUM 



81 



69 



Covacevich, 1997. From joint 
morphological/molecular com- 
parisons and field work in previously 
unexplored rainforests, we can now 
present a phylogeny (based on DNA 
sequences from the mitochondrial 
cytochrome b gene), erect a new 
genus to accommodate 'Saltuarius' 
occidtus and provide substantial new 
data (including descriptions of two 
new species) on Phyllurus spp. 

METHODS 



GENETICS. A 399 base pair 
fragment of the mitochondrial 
cytochrome b gene, corresponding to 
codons 1-133, was ampHfied and se- 
quenced from at least two individuals 
from each species of leaf-tailed gecko 
(Appendix 1 ). Protocols for amplifi- 
cation and sequencing follow those 
outlined in Couper et al. (1997). 
Sequences were aligned by eye using 
the translated amino acid sequences. 
For phylogenetic analyses, each 
nucleotide position was treated as a 
single character with up to four 
unordered states. The most parsimon- 
ious tree was found using equal- 
weights parsimony (all characters 
equally weighted and unordered), as 
well as a variety of weighting 
schemes. Additionally, maximum 
likelihood analyses, with a variety of 
models of nucleotide substitution 
were performed. In all analyses, 
sequences from Carphodaciyhis 
laevis Giinther, 1897 were used as an 
outgroup to root the tree. Bootstrap 
resampling and parsimony criteria were used to 
assess support for the recovered phylogeny. 
PAUP* 4.0b2 (Swofford, 1999) was used for all 
phylogenetic analyses. 

MORPHOMETRICS. All specimens examined 
are held in the Queensland Museum. Measure- 
ments were taken using Mitutoyo electronic 
callipers. Supralabials, infralabials and subdigital 
lamellae were counted on both sides of specimens 
examined. External morphological characters 
follow Covacevich (1975) and Couper et al. 
( 1 993). Skeletal definitions follow Bauer ( 1 990). 
Abbreviations: SVL, snout to vent length; T, tail 
length, from posterior margin of cloaca to tip of 
tail; TT, attenuated tip of original tail; HL, head 



100 



100 



m. 



S. salebrosus Blackdown 
S. salebrosus Bulburrin 
I S. cornutusAtherton 



S. cornutus MT 
— S. cornutus 8T 



100 



95 



3^ 



100 



63r 



100 



100 



S. cf. wyberba Chaelundi' 
. cf. wyberba Chaelundr 
S. wyberba Girraween 
S. wyberba Girraween 
S. wyberba Girraween 
S. swaini Lamington 
S. swaini MtTamborine 
Pnepthys Eungella 
Pneplhys Finch Hatton 
— Pchampionae sp. nov. 



Pisis 



97 



Possa Mt Dryander 
Possa Conway 
~ Possa MtOssa 



96 



Pplaturus Gosford NSW 
Pcaudlannulatus Bulburin 



Pcf. caudlannulatus Oakview 



100 



■ Pannnicola sp. nov. 
' Pamnicola sp. nov. 
Orraya (gen. nov) occultus 
C. laevis Southern Wet Tropics 
- C. laevis Northern Wet Tropics 



0.05 



Branch Length 

FIG. 1. Phylogeny of leaf-tailed geckos based on a 399 base pair 
fragment of MtDNA cytochrome b gene. Sequences from 
Carphodactylus laevis were used as an outgroup to root the tree and 
bootstraps >50% are presented (see Appendix 1 for specimen 
details). * Possible new species; here referred to Saltuarius cf. 
wyberba, pending collection of more specimens. 



length; HW, head width; S, snout length; LI, 
length of front leg, axilla to tip of longest digit; 
L2, length of hind leg, groin to tip of longest digit; 
NL, neck length, axilla to posterior margin of ear. 

PHYLOGENY. The mitochondrial cytochrome b 
DNA sequence data contained 183 parsimony- 
informative characters which provided good 
resolution of the phylogenetic relationships 
among species of leaf-tailed geckos. One 
hundred heuristic searches with random taxon 
addition and all characters unordered and equally 
weighted resulted in three equally parsimonious 
trees of 620 steps (CT = 0.463). The strict 
consensus of these trees, which differed only in 
the placement of P. platurus, is shown in Fig. 1. 
Bootstrap support for the clade containing all 



AUSTRALIAN LEAF-TAILED GECKOS 



255 



Saltuarius, except 'Saltimrius ' occultus (here 
recognised as generically distinct) is relatively 
high, while support for the monophyly of 
Phyllurns is somewhat weaker. Phylogenetic 
analyses and bootstrap resampHng with various 
w^eighting schemes to account for dilTerences 
among codon positions in rates of substitution 
and transition/transversion ratios resulted in the 
identical topology with similar bootstrap support 
to the equal weights bootstrap tree. Similarly, 
maximum likelihood analyses under a wide 
range of nucleotide substitution models (Jukes- 
Cantor, HKY85G, General time reversible) 
resulted in trees that did not differ from the equal 
weights parsimony tree. Species in the genus 
Phyllurus, exclusive of P. amnicola, are 
characterised by a derived karyotype as well as a 
number of morphological synapomorphies (see 
Phyllunis spp. nov.). Tlie karyotype of P anmicola 
is unknown but morphological synapomorphies 
are consistent with the DNA sequence data in 
supporting the inclusion of P. anmicola in the 
genus PhyUurus. 

Authorships for three sections of this paper 
(Orraya gen. nov., PhyUurus anmicola sp. nov. 
and PhyUurus champiouae sp. nov.) do not 
follow that of the paper as a whole. 

Orraya gen. nov. 
Couper, Covacevich, Schneider & Hoskin 
(Fig- 2) 

Saltuarius occultus Couper, Covacevich & Morilz. 1 993. 

A parsimony analysis of morphological and 
karyotypic characters performed by Couper et al. 
(1993) resulted in the recognition of two mono- 
pliyletic subgroups within the leaf-tailed geckos 
— P/7v/////-wi' Goldfliss, 1820 {P. caudianrndatus, 
P. is is, P. nepthys^ P. ossa and P. platufus) and 
Saltuarius Couper, Covacevich & Moritz,1993 
(S. cormitus, S. occultus, S. salebrosus and S. 
swaim). These clades were strongly supported in 
a bootstrap analysis present in 98% of 
pseudoreplicates. However, Couper et al. (1993) 
also observed evidence for grouping of 5*. 
swaini, S. conmtus and S. salebrosus to the 
exclusion of .S. occultus ..." and that Saltuarius 
occultus has skeletal characters (elongation of 
the cervical vertebrae and 3 lumbar vertebrae) 
that are unique amongst its congeners. Derived 
characters, including elongate cervical vertebrae, 
along with DNA sequences from the mito- 
chondrial cytochrome b gene (Fig. 1, Table 1), 
support the recognition of a new genus to 
accommodate 'Saltuarius ' occultus. In our 
phylogenetic analyses, this taxon is the sister 



group to the clade containing PhyUurus spp and 
Saltuarius spp, 

TYPE SPECIES. Orravaocculnts (Couper, Covacevich & 
Moritz, 1993). 

ETYMOLOGY. 'Orra>a' is the Morrobalama language 
word tor 'older brother' (Gunnawanu N. & KuUakulla, 
M., 1994) refering to the phylogenetic relationship 
between occultus and its PhyUurus and Saltuarius 
'siblijigs'. Peach Ck, the type locality of Orraya occultus 
drains the MclKvraith Ra. the traditional land of the 
MoiTobalarna. 

DIAGNOSIS. Orraya gen. nov. can be separated 
easily from all other Australian padless caipho- 
dactyline genera by the following combined 
apomorphies: cervical vertebrae greatly elongated; 
three lumbar vertebrae (defined as non-rib- 
bearing vertebrae, immediately anterior to 
sacrum); male preanal organs greatly enlarged; 
regrown tail with broad, spinose tubercules on 
margins. A detailed description of Orraya 
occultus (as Saltuarius occultus) was provided by 
Couper et al., 1993: 104-106. For details of 
original tail, see Lethbridge et al., 1994. 

PhyUurus spp. 

NEW SPECIES. The new species (from Mt 
Elliot, NEQ and Cameron Ck/Blue Mtn, MEQ) 
are assigned to PhyUurus by the following synapo- 
morphies: anterior-most autotomy septum in fifth 
caudal vertebra; no enlarged poslmental scales; 
tail finely- attenuated and tenninating in a small 
knob; ro.stral scale partially, or totally divided; 
males without preanal pores (polarity delemiined 
by Bauer, 1990). This assignment is supported 
fnrther by the following character states (after 
Couper et al., 1993): nostril not in contact with 
rostral scale; imterior margin of interclavicle with 
a distinct process; axilla invaginated; epipubic 
cartilage small to moderate and original tail 
simply flared. 

PhyUurus amnicola sp. nov. 
Hoskin, Couper, Sclineider & Covacevich 
(Fig. 3) 

ETYMOLOGY. From llie Latin 'dwelling by a river', a 
reference to the type locality. 

MATERIAL. HOLOTYPE: QN4J64408 ?, Alligator Ck, 
Mt Elliot, Bowling Green Bay NP (19°28'S, ]46°59'E) 
NEQ, C. Hoskin & J. Gratten, 1 Feb 1998. PARATYPES: 
QMJ64406-07, J67852 as for holotype. 

DIAGNOSIS. P anmicola can be separated from 
its congeners by the following combined char- 
acters: large size (maximum SVL >I10 mm); 



256 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 2. Orrava occidtiis (OMJ62596), Peach Ck, Mcliwraith Ra„ Cape York Peninsula (I3°45'S, 143°19'E) 
NEQ. (JeiT Wright) 



spinose body tubercules very small; leaf-shaped 
tail with anterior-most band broken but spanrring 
full tail width; rostral partially divided with 5-6 
scales along its dorsal margin. (Note: SVL 
includes three specimens measured by one of us 
(CJH) in the field (IS 104mm, ? 1 13mm). These 
measurements have been included in the 
diagnosis but not in the morphometries for the 
species description.) 

DESCRIPTION. SVL (mm): 90.3-103 (n = 4, 
mean = 96.5). Proportions as % SVL: LI 
43.9-45.1 (n = 4, mean - 44.6); L2 53.8-58.9 (n = 
4, mean = 56.2); T 86.1 (n = 1); TT36.9(n= 1); 
HL27.7-29.1 (n = 4,mean = 28.2);liW20.7-2L4 
(n = 4, mean = 2 1 . 1 ); S 1 2.4-1 3.2 (n = 4, mean = 
12.7); NL 20.7-23.3 (n = 4, mean = 21.6). 

Head large, depressed, triangular, distinct from 
neck; covered in small granules which are 
intermixed with larger conical tubercles; skin of 
head not co-ossified with skull; deep, vertical 
groove partially dividing rostral scale (n = 4); 
rostral excluded from nostril; 5-6 scales along the 
dorsal margin of rostral shield (n = 3); ear 
opening elliptical, vertical, much less than half as 
large as eye; supralabials 15-16 (n = 8, mean = 
15.5); infralabials 13-15 (n = 8, mean = 13.9). 
Neck broad. Body depressed, covered in small 
granules; dorsal granules intermixed with larger 
conical tubercles; tubercles ver>' small on back. 
Hanks and sides of neck; basal scales surrounding 
tlank tubercules not enlarged; no enlarged 
tubercules or granules on ventral surface of body. 
Preanal pores absent. Axilla moderately to 
deeply invaginated. Limbs long, covered in small 
pointed tubercles dorsally; lacking enlarged 
tubercules on ventral surface, except on upper 
forelimb; digits strong, strongly compressed 



distally; subdigital lamellae (fourth toe) 22-25 (n 
= 8, mean = 20.3). Original tail (n = 1 ) depressed, 
llared, contracted at base and attenuated at tip, 
terminating with a minute rounded knob; dorsal 
surface of flared portion with prominent enlarged 
spinose tubercules on basal l/3rd and along 
margins; 6 rows of minute spines across basal 
portion of attenuated tail-tip; attenuated tip 
accoimts for 42.9% of total tail length; ventral 
surface smooth with a slight depression along 
midline (excluding attenuated tip). Regenerated 
tail (n = 3) depressed, broad and strongly llared, 
contracted at base and attenuated at tip; covered 
with uniform granules, except on basal margin 
which has small spinose tubercules; ventral 
surface without groove along midline. 
Pattern. In spirit, dorsal base colour beige with 
irregular dark brown blotches on head, body and 
limbs; blotches on body tend to be aligned trans- 
versely. Limbs banded; digits strongly banded; 
inner anterior digit with reduced pigment. Body 
and limbs ventrally off-white to cream 
(immaculate); labials off-white, mottled with 
brown. Original tail dorsally tan/grey, marked 
with irregular dark brown blotches; six cream 
bands on tail, only those on attenuated portion (4) 
extending to ventral surface; ventrally cream, 
peppered with brown specks. Regenerated tail 
lacking cream bands; dorsally tan/grey, mottled 
with dark brown blotches; ventral surface 
similar, but with reduced pigmentation. 

Skeletal Features. Material examined: (radio- 
graphs) QM.T64406-08; (alizarin stained) 
QMJ67852. Supraocular portion of frontal 
grooved; anterior process of interclavicle 
pronounced; epipubic cartilage moderately 
expanded; presacral vertebrae 26; sacral 
vertebrae 2; lumbar vertebrae 2; first autotomy 



AUSTRALIAN LEAF-TAILED GECKOS 



257 




FIG. 3.P/w!ltirus umnicola sp, nov. (holotype, QIVU64408; QM photographic collection, ima£»e ref. N W982), Ml 
Elliot, NEQ. (Jeff Wright) 



number of scales along dorsal margin of rostral 
shield (5-6 vs 9-1 1 and 8-11, respectively). 

GENETICS. Phyllunts amnicola shows a 
21-27% sequence divergence from all other 
Phyllurus spp. for the cytochrome b portion of 
mtDNA (Table 1, Fig. 1). The phylogenetic 
position of/* amnicola as the sister group to the 
remaining Phyllurus is not strongly supported by 
the cytochrome /) data (Fig. 1 ), but morphological 
characters support its placement in a mono- 
phyletic Phyllurus. 

HABITAT & DISTRIBUTION. Mt Elliot, 
Bowling Green Bay NP, SOkins SE of Townsville 
(19°28'S, 146^S9^E) NEQ (Fig. 4). Found 
amongst boulders at 450m, in a moist 'tongue' of 
forest along a creek running fi^om the rainforested 
upper slopes of Mt Elliot (Fig. 5). Dense rain- 
forest is the dominant vegetation above 750m. A 
recent survey (CJH and J. Gratten, Oct., 1999) 
showed P. amnicola to be relatively common 
between 400- 1 000m along a major drainage line. 
Surveys on the summit of Mt Elliot have failed to 
locate this species. 

HABITS. Most specimens were found head 
down, on granite boulders close to pemianent 
water. Se\ eral were foraging on rocks directly 
above flowing water and one was observed 



septum on postsacral vertebra 5; abdominal 
vertebrae bearing reduced ribs 4; rib-lree cer\'i- 
cals 3; sternal ribs 3; mesostemal ribs 2. 

Holotype Data, QMj64408, 9 : SVL90.3mm; LI 
40.6mm; L2 53.2mm; T 77.8mm; TT 33.4mm; 
FIL 25. 1mm; HW 18.8mm; S 1 1. 3mm; NL 19.0mm; 
supralabials 15/16; infralabials 15/14; subdigital 
lamellae 22/22. 

COMPARISON. Phyllurus amnicola can be 
confused with only its congeners. P. amnicola is 
distinguished from P caudiannulatus by tail 
shape (flared vs cylindrical); from P. platurus by 
pattern of original tail (flared portion with 
distinct white bands vs without white bands); 
from P. is is by pattern of original tail 
(anterior-most band spanning full width of tail vs 
anterior band reduced, with two narrowly-spaced 
midline blotches); from P. nepthys by colour/ 
pattern of ventral surface (immaculate vs pep- 
pered with brown specks); from P. chcunpionac 
by spinosity of original tail (enlarged spinose 
tubercules restricted to anterior third of tail and 
tail margins vs tail covered with spinose 
tubercules) and from P. ossa by the rostral 
groove/s (one groove partially dividing rostral vs 
1-3 grooves, usually 3, only rarely 1 or 2, 
partially dividmg rostral). P. amnicola is further 
distinguished from P. isis and P. ossa by the 



258 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Townsville 




Rockhampton 



! Brisbane 



NSW 



Sydney 



FIG. 4. Occurrence of Phyllwus spp. in eastern 
Australia; \=P. amnicola, 2 =P. ossa, 3 = P. nepthys, 
4 = P. isis, 5 = P. championae, 6 = P. cavdiamndatus, 
1 ^ P. platurus. 

foraging in a thin film of water in the splash zone 
of a cascade. All specimens, but one, have been 
found on rocks. The exception was on a thin tree 
trunk amongst boulders. Activity began soon 
after dark, even during persistent rain. Of 27 
specimens (14c?'s, II 9 's and 2 juveniles) en- 
countered during two nights (Oct., 1999), 70% 
had regenerated tails. This proportion was similar 
in both sexes. 

REPRODUCTION. One gravid female, captured 
and released (4 Feb., 1998), contained a single, 
shelled egg. A male (QMJ67852) that died in 
captivity (Dec, 1 998) was sexually mature, with 




FIG. 5. Alligator Ck, Mt Elliot, NEQ, the type locality 
for P. amnicola sp. nov. (Conrad Hoskin) 

sperm present in its epididymis (inferred by 
opacity). Fourteen mature females were en- 
countered on the first two nights of Oct., 1999. 
Eight of these carried well-developed, shelled 
eggs (3 with I egg, 4 with 2 eggs, 1 with 3 eggs). 

CONSERVATION. P. amnicola is one of the 
most narrowly restricted reptile species in 
Queensland. However, it is well protected. The 
only known locality for this species is in Bowling 
Green Bay NP which is not subject to any known 
threatening processes. The potential effect of fire 
on pockets of riparian rainforest at and near the 
type locality is not known. 

Phyllurus championae sp. nov. 
Schneider, Couper, Hoskin & Covacevich 
(Fig. 6) 

ETYMOLOGY. Named for Irene Champion, a Resource 
Ranger with Queensland Parks and Wildlife Service, 
Mackay, MEQ, who focussed the attention of one of us 
(CJS) on Cameron Ck/Black Mtn, as a phytogeo- 
graphically interesting area, possibly pointing to the 
presence of unusual fauna. 



AUSTRALIAN LEAF-TAILED GECKOS 259 




FIG. 6. Fhylhirns championae sp. nov. (QM photomaphic collection, imauc ici. NX/."^8) Cameron Ck, MEQ- 
(Jeff Wright) 



MATERIAL. HOLOTYPE; QMJ64847 Cameron Ck, 
6.5km WNWKoumala(21°34'24"S,149°ll'06"E) MEQ, 
P. Couper & C. Hoskin, 18 April 1998. PARATYPES: 
QMJ62757-58, J62766, J63907. J64845-46, J64848, 
locality as for holotype; J64854-64 Blue Mtn (2r36^S, 
]48°58^E)MEQ. 

DIAGNOSIS. P. championae almost invariably 
(18/19) has a Hilly divided rostral scale. This 
feature, combined with a leaf-shaped tail, 
distinguish it from all other Phvlliirus spp. A 
specimen of P. championae with an only partially 
divided rostral could be confused with some 
specimens of P. ossa which have a rostral 
partially divided by a single groove. (This is a 
rare stale in P. ossa which usually has 2 or 3 
partial grooves). Erom such specimens ofP ossa, 
'partial single groove' specimens off! championae 
can be distinguished readily by a straight groove 
vs a Y-shaped groove. 

DESCRIPTION. SVL(mm): 33.3-80.6 (n = 19, 
mean = 61.5). Proportions as % SVL: LI 41.2- 
48.0 (n - 1 9, mean = 44.0); L2 52.5-60.5 (n = 1 9, 
mean = 56.50); T 64.2-8 1 .7 (n = 8, mean = 75.7); 
TT 28.9-40.7 (n - 8, mean = 35.7); HL 28.9-3 1 .8 
(n - 19, mean - 30.1 ); HW 23.1-26.4 (n - 19, 
mean = 24.40);S I2.4-I4.6(n=19,mean=13.1): 
NL 18.8-23.6 (n= 19, mean = 20.6). 



Head large, depressed, triangular, distinct from 
neck; covered in small granules which are inter- 
mixed with larger, conical tubercles, extremely 
prominent on snout; skin of head co-ossified with 
skull; deep, vertical groove totally dividing 
rostral scale (n=18) or (rarely) partially dividing 
rostral scale (n=I ); rostral excluded from nostril; 
5-8 scales along dorsal margin of rostral shield (n 
= 10); ear opening elliptical, vertical, much less 
than half as large as eye: supralabials 12-14 
(n=38, mean^l3.3, mode==i4); infralabials 1 1-15 
(n^38, mean=12.7, mode^l3). Neck broad. 
Body depressed, covered in small granules; 
dorsal granules intennixed with larger, conical 
tubercles; tubercles small on back, pronounced 
on flanks, most prominent on sides of neck; basal 
scales surrounding flank tubercules only slightly 
enlarged; no enlarged tubercules or granules on 
ventral surface of body. Preanal pores absent. 
Axilla deeply invaginated. Limbs long, covered 
in large pointed tubercles dorsally; without 
enlarged tubercules on ventral surface, except on 
upper forelimb; digits strong, compressed distal- 
ly; subdigital lamellae (fourth toe) 16-20 (n = 38, 
mean = 17.9, mode ^ 17). Original tail (n = 8) 
depressed, flared to carrol-shaped, contracted at 
base and attenuated at tip, terminating with a 



260 



MLMUJRS OK i HE QUEENSLAND MUSEUM 




FIG 7. A-, t ameron Ck, MtQ, the type locality for/* vhumpioime sp. nov. B, dry rainforest below liie summit of 
Blu&Mtn,JVlEQ^theBecondktv)wnJocaLity'ror/! champiai^iae^i^^t^i^. (Conrad Hosldn) 



minute touod-dd knob; ciov^d dorsally witft 
tUltncrous modcnitc-si/cd. spinnsc tubcrculcs 
Which become smaller along the \eriebral line, 
tubercles terminate approximately half-way 
aloi^lhc attenuated tip which is lpng;-Srows.of 
twlargcd spines across the basal iJorS<5tv 
attenuated tail-tip; attenuated tip accounts for 
38-52% of total tail length; ventral surface 
smooth, or with slight depression along midline. 
Regenerated tail: m - 10) depressed, broad and 
strongly flared, contracted at base and attenuated 
at tip; with small, spinose lubercules which are 
tftost prominent around the edges; ventral surface 
u itluuit uroove along midline. The Cameron Ck 
specimens are signilleaiilly smaller than those 
Irom Blue Mtn (max SVL - O'-^mm, ii - 7, small 
juveniles excluded, mean ^ Ol.ynini vs max SVL 
8 1 mm, n - 9, snial 1 juveniles occluded, mean = 
69.6mm; Student's t-Test,. t, ij - 2.14, 0.05> 
P>0.02S). 

Paitenh In spirit, dorsal base coioiu: mid brown 
witli icregular, black blotches oti lufatl* body ail4 



limbs'. tJi^ife obscurely Bande^ iwner antferibr 

les digit not significonily lighter thfiUlOflacrs, P^t'dy 
and limbs \cntrally otV-white^!& Cfcam; labials 
otT-white, moitlcd with ItfOs^* Original tail 
dorsally tan. heavily mottled "Wifli black (ahnost 
entirely black in hatchlings5; liv« white bands on 
laiE only those (2-3) on attenuated portion ex- 
tending to ventral surface; ventral ly cream with 
grey mottling. Regenerated tail lacking cream 
bands; dorsally, tan/'grey with black blotches; 
ventral surface simi&, mt with reduced pig^ 
mentation. 



Shst&tqlF&aHtm- M^l^^ examined; (mUpgraphsJ 

J64864;Xa!izarin .^ed) QM.164863. Stipra- 
oeular portion of frontal flat; anterior process ol' 
interela\ ieie prtmounced; epipubic carlilauc not 
expanded: presacral vertebrae 26; sacral 
vertebrae 2; lumbar vertebrae 2; first autotomy 
septum on postsacral vertebra 5; abdominal 
vertebrae bearing reduced ribs 4; rib-free 
wrvieals 3; s<emql ribs 2; mesostemal libs 3. 



AUSTRALIAN LEAF-TAILED GECKOS 



261 



TABLE L Kimura 2-parameter distance estimates (Kimura, 1980) between species and populations within 

spec ies for 3 99bp cytochrome h sequence data. S pec ies are numbered In the same order across the top of the data 
matrix. See Appendix 1 for specimen details. 

1 2 3 4 5 6 7 8 9 10 11 12 13 14 

1 S. salebrosus Blackdown 

2 S. salebrosus Bulburrin 0.0625 

3 S, cornutus Atherton 1833 0.1760 

4 S. cornutus MalbonThompso 0.1872 0,1722 0.0076 

5 S. cornutus BigTableland 2039 0.1785 0.0894 0960 

6 S. cf. wyberba Chaelundi 1987 0.2119 0.2677 0.2726 2733 

7 S, cf wyberba Chaelundi 0.1987 0.2119 02677 2726 2733 0.0054 

8 S. wyberba Girraween 0,2280 0.2376 0,2640 0.2687 0.2921 0.1321 1321 

9 S. wyberba Girraween 0,2244 0,2384 2602 0,2648 0,2878 0.1288 0,1288 0025 

10 S. wyberba Girraween 0.2208 0.2347 0,2563 2610 0,2835 0.1255 0,1255 0.0050 0025 

11 S. swaini Lamington 0.2251 0.2087 0,2296 0,2383 0,2627 0.1520 1600 0,1733 0,1700 0,1668 

12 S, swaini MITamborine 0.2244 02081 0.2260 0.2346 0.2585 0,1514 0.1594 0.1727 0,1694 0.1661 0,0050 

13 P, nepthys Eungelia 0.2980 0,3093 0,2862 0,2862 0,2756 0,2514 0.2610 0,2889 0.2849 0,2810 0,2602 0.2594 

14 P, nepthys Finch Hatlon 2929 0,3041 0,2813 2813 0,2806 0,2561 0,2658 2840 0,2800 0,2761 0,2555 0.2548 0025 

15 P isis 03051 0.2911 0,2966 3065 2973 0.2591 2686 2797 0.2757 0.2718 2642 0.2591 0,1544 1581 

16 P, ossa MtDryander 2765 0,2679 0,2909 3007 0,2774 0,2453 0.2546 0.2869 0.2876 0,2836 2591 2585 0,1512 0.1549 

17 P ossaConway 2757 2765 0,2966 0.3065 2884 0,2457 02551 0.2925 0.2933 0,2893 0,2642 0,2636 1516 0,1553 

18 P ossaMtOssa 3020 0,2831 0.3033 0.3134 0.2748 0.2719 0,2719 0.3042 3051 0.3010 0.2992 02983 0.1416 1452 

19 P championae sp. nov. 3001 0,3010 0,3218 0.3271 03136 0.2685 0,2786 2942 02951 0,2911 2893 0.2885 0,1563 0.1600 

20 P piaturus 0,2510 0,2752 0,2496 0,2586 0,2702 0,2499 0,2596 2414 0.2376 0.2339 0.2518 0,2510 0.1936 0,1975 

21 P caudiannulatus 0,2610 0,2853 0.2829 0,2925 0.3130 0,2650 0,2749 0,2789 0,2797 0,2757 0.2983 0,3024 0,2405 0,2450 

22 P. cf, caudiannulatus Oakvie 0,2742 0,3010 0.2558 0,2649 0.3094 0,2699 0,2799 0.2781 0,2742 0,2702 0.2642 0,2681 0.2056 0,2097 

23 P, amnlcola sp, nov, 0,2579 2496 0,2688 0,2734 0,2859 0,2563 0,2470 0,2514 0,2476 0.2438 0,2558 0,2508 0.2420 0,2376 

24 P amnicola sp nov, 0,2534 0.2451 0,2688 0,2734 0.2861 0,2516 2424 0,2470 0,2432 0,2395 0,2514 0,25Q8 0.2376 0.2333 

25 Orraya(gen. nov,)occultus 0,2632 0,2640 0.2309 0,2351 2575 0,2336 0,2336 2565 0,2572 0,2534 0.2489 0,2482 0.2427 0.2383 

26 C. laevis BigTableland 3033 0,2893 0,2448 0,2491 0.2759 0,2702 0,2801 0,2880 0,2840 0.2880 0,2656 2649 0.2521 0,2568 

27 C laevis Alherton 3074 0.3033 0.2530 2574 0.2803 0,2845 2948 0,3020 2980 0.3020 0,2558 0,2642 0.2559 2606 







15 


16 


17 


18 


19 


20 


21 


22 


23 


24 


25 


15 


P. rsis 
























16 


P. ossa MtDryander 


1230 






















17 
18 


P ossa Conway 
P, ossa MtOssa 


1235 
0,1306 


0.0206 
0,0595 


0,0568 


















19 


P, championae sp, nov. 


0.1682 


0.1727 


0.1694 


0,1787 
















20 


P, piaturus 


2000 


0,1857 


0,1853 


0,1941 


1970 














21 


P, caudiannulatus 


0.2383 


0.2145 


0.2194 


0,2280 


0,2223 


0, 1 887 












22 


P, cf caudiannulatus Oakvie 0,2333 


0,2211 


0.2217 


0,2145 


0.2390 


0.1853 


1452 










23 


P amnicola sp. nov. 


2097 


0,2450 


0.2488 


2442 


0.2310 


2123 


0,2451 


02687 








24 


P. amnicola sp, nov. 


0,2139 


2405 


0.2442 


2398 


2266 


2081 


0,2407 


2640 


0025 






25 


Orraya (gen, nov,) occultus 


0.2290 


0.2046 


0.1931 


1970 


0.2555 


0.2122 


0,2563 


0,2451 


0,2314 


02272 




26 


C, laevis BigTableland 


0,2555 


0,2656 


0.2713 


2829 


0,2450 


0.2390 


2831 


0,2951 


2625 


0,2579 


0.2092 


27 


C, laevis Atherton 


0,2782 


0.2791 


0,2849 


2970 


0,2580 


0,2383 


0,2870 


0,2749 


2773 


0,2726 


0,2127 



Ho/otvpe Data. QMJ64847, S; SVL 59.28mm; 
LI 25.57mm; L2 32.80mm; T 48.13mm; TT 
24.L3mm; HL 17.81mm; HW 14.20mm; S 
7.77mm; NL lL34mm; supralabials 13/14; 
infralabials 13/13; subdigilal lamellae 18/17. 

COMPARISON. R championae can be confused 
with only its congeners. P. championae is 
distinguished from P. caudiannulatus by tail 
shape (leaf-like vs cylindrical); fi*om P. piaturus 
by colour pattern of anterior portion of original 
tail (with white bands vs without white bands); 
from P amnicolahy spinosity of original tail (tail 
covered with spinose tubercules vs enlarged 
spinose tubercules restricted to anterior l/3rd of 
tail and tail margins); from P. isis by spinose 
Hank tubercules (pronounced vs very small); 
from P nepthys by colour/pattern of ventral 
surface (plain vs peppered with brown specks) 



and from P ossa by rostral groove/s (usually one 
groove dividing rostral, or a straight vertical 
groove partially dividing rostral vs 1-3 grooves, 
usually 3, only rarely 1 or 2, partially dividing 
rostral). 

GENETICS. P championae shows 16-24% 
sequence divergence from all other PhyUurus 
spp. for the cytochrome h portion of mtDNA 
(Table 1). The Cameron Ck and Blue Mtn 
populations showed no within, or between, 
population polymorphism. Phylogenetic 
analyses tirmly place P. championae within the 
monophyletic MEQ species group of PhyUurus 
(P. isis. P. nepthys and/^ ossa. Fig. 1), though its 
precise relationship to these taxa is not well 
resolved. 



262 



MEMOIRS OF THE QUEENSLAND MUSEUM 



HABITAT & DISTRIBUTION. P, championae is 
known from only two localities, Cameron Ck 
(2r34'24"S, 149°ir06"E) and (2 1 km to the 
west) Blue Mtn(21°36'S, 148°58'E) MEQ, (Fig. 
4). The type specimens were collected at ahitudes 
between 200m (Cameron Ck) and 700m (Blue 
Mln) in notophyll rainforest/microphyll 
rainforest. 

HABITS. All specimens have been on rocks or on 
the trunks of trees near rocks. The Cameron Ck 
(2r34'24"S, 149^11 ^06"E) specimens were 
collected on the edges of a permanent creek on 
the eastern side of Black Mtn (Fig. 7a). Blue Mtn 
(2r36'S, 148°58'E) specimens were active on a 
scree slope, in dry rainforest, just below the 
summit (Fig. 7b). Activity began soon after dark. 

REPRODUCTION. Gravid females were present 
in the Cameron Ck population during Dec, 
1996/Jan., 1997. A female measuring 68.7mm 
SVL and weighing 6.2g (QMJ62757^ laid two 
oval-shaped eggs on 6 Jan., 1997. These 
measured 18.35 x'9.0 1mm and 17.72 x 8.90mm 
and weighed l.Og and 0.9g, respectively. The 
relative clutch mass (RCM (1) after Greer, 1989) 
equalled 30.6%. This corresponds closely with 
that ofP.platunis (29%, Greer, 1989). 

A male (QMJ64863, SVL 72.3mm) from Blue 
Mtn, collected on 19 Apr., 1998, was in peak 
reproductive condition with sperm present in its 
epididymis (inferred by opacity) and a turgid 
testis. From this we infer that mating may occur 
in autumn and that females store sperm tlirough 
winter. A similar reproductive strategy has been 
suggested for platiirus (Greer, 1989). 

CONSERVATION. P. championae appears to be 
common at both localities from which it is 
knowii. Suitable similar habitat in adjacent areas 
may support this species. Blue Mtn is freehold 
and thus, potentially, could be cleared. However, 
given that the known leaf-tail site is on an 
elevated scree slope, the chance that this area 
would be tlirther disturbed by stock or humans 
seems remote. Cameron Ck, the type locality, is 
in State Forest and may be subject to timber 
harvesting. 

Phyllurus caudiannulatus 

The discovery of specimens treated tentatively 
as P caudianmdatits (QMJ62817, J63849-53, 
J63857) at OakN'iew SF (26°07'23"S, 1 52°l 9'0l"E), 
SEQ extends the range of this species 195km 
south of the only other known population 
(Bulburin SF, 24°3rs, 151°29'E), SEQ. 



Morphological ditTerences between the Oakview 
and Bulburin populations cannot be fully 
assessed presently, due to small sample size. 
Surveys in the intervening areas should help 
assess the significance of these difterences. For 
conservation purposes, the Oakview population 
of 'P. caudiannulatus' must be recognised as a 
genetically distinct unit (Fig. 1 ), given its geographic 
and genetic isolation and its uncertain taxonomic 
status. 

KEY TO AUSTRALIAN LEAF- TAILED 



GECKOS 

i .Nostril in contact with rosti'al scale 2 

Nostril not in contact with rostral scale 6 

2. Neck extremely elongate and slender . . . . O. occultus 

Neck broad 3 

3. Throat strongly tuberculate S. salebrosus 

Throat smooth 4 

4. ! .ower flank tubercules hooked and surrounded by spinose, 

basal scutes S. cormUus 

Lower tlank tubercules not as above 5 



5. Original tail bluntly tipped with large tubercules 

S. smiini 

Original tail finely tipped with only minute tubercules 
S. wyherha 

6. Tail cylindrical P. caudiannulatus 

Not as above (leal-shaped). 7 

7. Anterior ponion oforiginal tail without white crossbands 

or blotches P. planma- 

Nolasabove(vvith whitecrossbands) 8 

8. Venterdistinctly 'peppered' with brown . . . P.nepthys 

Notasabove(plain) 9 

9. Rostral scale completely divided P. championae 

Not as above (partially divided) 10 

10. Rostral scale partially divided by 2 or 3 grooves, 
occasionally by a single Y-shaped groove . . . P.ossa 

Nolasabove(partially dividedby a straight groove). . 1 1 

1 ! . Anterior flared portion oforiginal tail uniformly covered 
with enlarged spinose tubercules . . . . P. championae 

Not as above 12 

12. Fail predominantly black, with distinct white blotches 
present on either side of the vertebra! line .... P.isis 

Not as above (tail predominantly tan/grey, with anterior- 
most bands broken, but spanning tail width) 
P. amnicota 

DISCUSSION 

Twelve species of leaf-tailed geckos in three 
genera occur in eastern Australia (I3°45'S - 
33°53'S). Nine are obligatory rainforest/adjacenl 
wet sclerophyll forest species: Mcllwraith Ra., 
NEQ, Orrava occultus (Couper, Covacevich & 
Moritz, 1993); Wet Tropics (Big Tableland - 
Paluma), NEQ, .S". cormitus (Ogilby, 1892); Mt 
Elliot, NEQ, Phyllurus amnicola Hoskin, 
Couper, Schneider & Covacevich, 2000; Mt 



AUSTRALIAN LEAF-TAILED GECKOS 



263 



Dryander - Mt Ossa, MEQ, P. ossa Couper, 
Covacevich & Morilz, 1993; Mt Blackwood and 
Mt Jukes, MEQ, P. isis Couper, Covacevich & 
Morilz 1993; Clarke Ra. (Mt David - Credilon), 
MEQ, P. nepthys Couper, Covacevich & Moritz, 
1993; Black Mountain and Blue Mountain, 
MEQ, P. championae Schneider, Couper, Moskin 
& Covacevich, 2000; Many Peaks Ra. and 
Oakview Stale Forest, SEQ, P. caudiammlatus 
Covacevich, 1975; Great Dividing Range and 
foothills and Border Ranges (Mt Tamborine - 
Buladelah), SEQ-MENSW, S. swaiui Wells & 
Wellington, 1985. Two species are conlined to 
heaths associated with either sandstone or 
granites: Girraween National Park, SEQ, S. wy- 
herha Couper, Schneider & Covacevich, 1997; 
and Hawkesbur}' R. region, MENSW, P. platums 
(Shaw, 1 790). (3ne species, S. salehrosus, occurs 
amongst sandstone in open forests: Blackdown 
Tableland - Cracow, MEQ. It is also found in 
rainforest in the Many Peaks Range (24°3rS, 
l5r29'E), where it is sympatric with P. caudi- 
unnulalus. This is the only known area to support 
more than one species of leaf-tail. 

Tlie genetic analyses (Fig. 1 , Table 1 ) show that 
the depth of divergence amongst leaf-tails is high 
and that more-than-trivial differences between 
some taxa/populations remain to be clarified. 
Relative rates tests show that substitution rates 
among lineages do not differ significantly 
suggesting that a molecular clock may apply. By 
using the break-up of Gondwanaland to estimate 
substitution rates in this portion of cytochrome b 
among carphodact>'line geckos Irom Australia, 
New Zealand and New Caledonia (Schneider, 
unpublished) it is estimated that the Kimura 
two-parameter distance between two lineages 
accrues at a rate of ca. 0.0042 ± 0.0002 per 
million years (0,0021 per lineage per million 
years). Using this calibration, the split between 
Saltuariiis sensu stricto and Phyllurus is ca. 
58-74mya. The divergence among species in 
MEQ is ca. 3 1 -38mya, and this is nearly identical 
to the estimated time of divergence between the 
Oakview and Bulburin populations of P. coudi- 
anmilatus. Estimating genetic distances among 
taxa using only transversions (which are more 
likely to accrue linearly with time) does not sub- 
stantially change these time estimates. \Salfuanus 
wvberha' populations from Chaelundi SF, NSW 
(30°0r07"S, 152°30^02^^E) and Girraween, 
SEQ (28°50'S, 151°56.04E, the type locality) 
dilTcr to the same decree as P. isis (from Mt 
Blackwood, 2 r02'S, r48°56'E) and P ossa {Mt 
Ossa, 20°56'S, I48°49'E, only 14km to the 



north). However, more specimens must be 
examined before the status of the Chaelundi 
Saltuariiis specimens can be determined. 

Recognition of Orraya occultus as distinct 
from the other large leaf-tails (Salfiiarius spp.) is 
significant given the general paucity of 
endemism in the reptiles of the Mcllwraith Ra. 
rainforest isolate. (Couper et al., 1993 observed 
this in relation to the Wet Tropics rainforests 
where 2/3 of the rainforest reptile species are 
endemic to the area). Orraya is the sole terrestrial 
vertebrate genus not represented in rainforest 
isolates further south. 

The discovery of P awtiicola on Mt Elliot, 
NEQ is noteworthy for two reasons. First, recog- 
nition of this species brings to two the number of 
vertebrate species endemic to Ml Elliot (the other 
species is the microhylid frog, Cophixaliis 
mcdonaldi Zweifel, 1985), highlighting the 
evolutionary uniqueness ol'this rainforest isolate. 
Second, il extends the range Phyllurus some 
200km NW of its previously-known, northern 
limit of occurrence (Mt Dryander, 20''15'S, 
148°33'E, MEQ), across what has been tenned 
the 'Burdekin Gap' (Joseph el al., 1993). This 
expanse of dry woodland between Mt Elliot 
( 1 9°30'S, 146°58'E), NEQ and Bowen (20'^0 1 'S, 
148° 1 5'E), MEQ has separated the faunas of two 
major zoogeographic regions, the Wet Tropics 
and Central Mackay Coast, for an 'evolulionarily 
long period' (Joseph el al., 1 993 ). There is a deep 
divergence between P anmicola and its congeners 
immediately south of the Burdekin Gap (ca. 50-60 
mya). The mlDNA sequence divergence between 
P, amnlcola on the one hand, and P. championae, 
P. isis, P. nepthys, P. ossa, on the other, is thus 
nearly as great as that belM'een these species and 
those of Saltuarius spp. With the recognition of P. 
anmicola, there is now an overlap in the max S VL 
between the largest member of Phyllurus (P. 
amnicola, max SVL = 1 1 3mm) and the smallest 
species of Saltuarius {S. wyherha, max SVL = 
109mm). 

Description of P. championae brings to four the 
number of Phyllurus species known to be 
confined to rainforests of the Central Mackay 
Coast Biogeographic Region, MEQ. Couper et 
al. (1993) commented on aspects of the 
zoogeography of the other species confined to 
this area, all within 100km of each other (/! isis, P 
nepthys and P. ossa). It is now clear that the 
species of leaf-tailed geckos in rainforests of 
MEQ represent the relictual distribution of an 
ancient group. It has been thought that 



264 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Pleistocene ice age contraction of rainforest 
resulted in speciation among rainforest 
endemics, but the deep genetic divergence among 
leaf-tailed geckos precludes any role for 
Pleistocene speciation. 

Whether P. caudiannulatus (sensu stricto) 
occurs between Bulburin and Oakview State 
Forests, SEQ or is confined to the northernmost 
locality, remains to be ascertained. If the latter is 
the case, the Oak\'iew 'P. caudiannulatus' may 
represent another new species, pointing to a 
replication of patterns of relictual isolation 
already observed in MEQ. 

Most leaf-tails (Phyllurus spp., Saltuarius spp. 
and Orraya occultus) are very narrowly dis- 
tributed. Several {Orraya occultus, P. amnicola, 
P. isis and P. nepthys) are known from only single 
localities. Four other species (S. cormitus, S. 
swaini, S. wyberba and P. caudiannulatus) are 
narrowly distributed, being contined to small 
rainforest or rainforest and heath blocks. Under 
rUCN (1994) criteria, notwithstanding narrow 
distributions, all species should be categorised 
'Data Deficient'. Higher lUCN categories, which 
reflect concerns based on such single localit>'/ 
narrow ranges are all tied to knowledge of 
declines in populations and/or potential threaten- 
ing processes. Virtually all known leaf-tail 
localities are in state conser\'ation or timber 
reserves where, generally speaking, threats are 
presently low. Impacts of possible future timber 
harvest in some areas are not known. However, 
most of the knowTi, still healthy leaf-lail localities 
have, in the past, already been selectively logged, 
some extensively. 

ACKNOWLEDGEMENTS 

This work was supported financially by the 
Queensland Museum, Co-operative Research 
Centre for Tropical Rainforest Ecology and 
Management (University of Queensland), Centre 
for Conservation Biology^ {University of Queens- 
land) and the Forest Ecosystem Research and 
Assessment Unit of the Department of Natural 
Resources (Comprehensive Regional Assessment 
Project). We are grateftil for assistance in the 
preparation of this paper from Robert Adlard, 
Gary Cranitch, Ben Hamley, Luke Hogan, John 
Hooper, Michael Koch, Jacob Gratten, Heather 
Janetzki, Keith McDonald, Edward Meyer, Geoff 
Monteith, Craig Moritz, Geoff Smith, Joanna 
Sumner and Steve Wilson. Andrew Amey and 
Lauren Keim helped to improve the key. 



LITERATURE CITED 

BAUER, A.M. 1990. Phylogenelic systematics and 
biogeography of the Carphodactylini (Reptilia: 
Gekkonidae). Bonner Zoologische Mono- 
araphien 30: 1-2 17. 

COUPER. P.J., COVACEVICH, ,I.A. & MORITZ, C. 
1993. A review of the leaf-tailed geckos endemic 
to eastern Australia: a new genus, four new 
species, and other new data. Memoirs of the 
Queensland Museum 34(1): 95-124. 

COUPER, PJ., SCHNEIDER, C.J. & COVACEVICH, 
J. A. 1 997. A new species of Saltuarius (Lacertilia: 
Gekkonidae) from granite-based, open forests of 
eastern Australia. Memoirs of the Queensland 
Museum 42(1): 91-96. 

COVACEVICH, J. 1975. A review of the genus 
P/iyllurus (Lacertilia: Gekkonidae). Memoirs of 
the Queensland Museum 17(2): 293-303. 

GREER, A.E. 1989. The biology and evolution of 
Australian lizards. (Surrey Beatty & Sons: 
Chipping Norton, Sydney). 

GUNNAWARRA, N. & KULLAKULLA, M. 1994. 
Morrobalama- English. In Ogilvie, S. (compiler) . 
A wordlist of the Morrobalama (Umbuygamu) 
language of Cape York, Australia. (Umagico 
Council: Umagico). 

lUCN SPECIES SURVIVAL COMMISSION, 1994. 
lUCN Red List Categories. Prepared by the lUCN 
Species Survival Commission as approved by the 
40th meetina of the lUCN Council, Gland, 
Switzerland/30 Nov.. 1994. (lUCN Council: 
Gland). 

KIMURA, M. 1980. A simple method for estimating 
evolutionary rate of base substitutions through 
comparative studies of nucleotide sequences. 
Journal of Molecular Evolution 16: 111-120. 

LETHBRIDGE, RJ.. HAWKES. LA., ANTHONY, M. 
& MCGREGOR, M. 1994. New data on 
Saltuarius occultus, a recently described, poorly 
known leaf-tailed gecko. Memoirs of the 
Queensland Museum 37(1): 194. 

SWOFFORD, D.L. 1999. PAUP*. Phylogenetic 
Analysis Using Parsimony (* and other methods). 
Version 4. (Sinauer Associates Inc: Sunderland, 
Massachusetts). 

APPENDIX 1 

SPECIMENS EXAMINED. The following have 
been examined in addition to other material cited 
in Couper et al., 1993. 
Morphology: 

/^ caudiatmulatus - QMJ15619 (holoiype), J33684-86, 

J33706, J33709, J62817, J63849-53, J63857 
P. isis - QMJ535n (holotype), J53485-86, J53518. J53480, 

J53591,J53602-3 
P- nepthys - QMJ34058 (holot\pe), J34057, J351 14, J5703I, 

J655i 1, J65575, J65578, J65580, J65582, J65584, J65674 
P. ossa - QMJ53444 (holorvpe), J53389, J53392, J53426, 

J53428, J53443, J53445, J53447, 156311, J56773, J56791 
P plalurus - QMJ160, J31978, J56880-8 1 , ,156895 



AUSTRALIAN LEAF-TAILED GECKOS 



265 



Orrava ocxultus - QMJ37040 (holO^), H703t, J37038-9 

J60717, J62596 
a /«evisr. QMBI004. J65410'U, J65419 

Genetics: 

Phvllurus ammcolu - liver samples from QMJ64406-7 (Ml 

Elliot - 19°28'S. 146^59^E). 
P. caudiarmulatus - (Bulburin SF - 24°31'S, 15r29'E) liver 

Samjle firanQMJSUOa, 
P, cmdtanmtatus {Oakview - 26^0r25"S, l52'*19^0rE) - 

Hver sample from QMJ62817. 
P. championae - liver samples from QM.162757-58, J62766, 

J63907 (Cameron Ck - 2r34'24"S, 149^1 r06"E) and 4 

liver samples from the following series:- .164854, .164857, 

J64859, J64861-62 (Blue Mm - 2r36'S, 148'^58'E). 
P. to -tail tips from Mt Blackwood ( 21"02'S, 148"56"E). 
P. nepthvs - liver samples from QM.I5 1 1 1 & .15 1 098 ( Finch 

Hatt6nNP-21'^'06'S, 148"38"E). 
P, ossa - tail lips from Ml Ossa (20°56'S, 148°49'E). Brandy 

Ck (20°2rS, 148°4rE) and Mt Dryandar {2Q°15'S, 

148°33'E). 



P. pJatunts - liver samples fiom QMJ36S80-1 (via GosfM ^ 

33°24'S. 13I"21'E). 
SaUuarius lonmtus - liver samples from QMJ5 1 632 (Malbon 

Thompson Ra.- 17°07'S, 145^54'E) and tail tips from 

Athcrton (17''16'S, HS'^S^^'E) and Big Tableland ( 

15^'43'S, 145n7'E). 
S.. sdebrpBm ^ liv^ sao^l^ ^om (Bj^doSawn 

Tabletod-me^S, 14§°06'E) aud JS1090 »ulburin SF- 

24^31% l5l^m 
S. smbii - Hv^ SsirftpieS frtiffl QMT5 1^0 ( tatriington KP- 

28°14'S, 153°08'E) and .T51095 (Mt Tamborine - 

27°58'S, 153^irE). 
.S, wvherba - liver sample from QMJ5 1633 and tail tips (Gir- 

raweenNP- 28^^50' S, 151°55'E) 
S. ct: wvherba - (Chaelundi SF population - 30°0r07"S, 

15230^02"E & 30°O3'04"S, 152°2r36"E) tissues from 

AMR14 1964-5. 
Orrava occiiltus - tissues from QMJ60717, J62596 

(Mcllwraitli Ra. - 13°45'S, 143°19'E). 
Carphodactylus laevis - tails from Big Tableland (15°43'S, 

145°17'E) andMt Bartle Frere(17°24'S, 145M9'E). 



266 



MEMOIRS OF THE QUEENSLAND MUSEUM 



THE SCINCID LIZARD EGERMA MCPHEEI WELLS 
& WELLINGTON, 1984 IN QUEENSLAND. Memoirs of 
ihe Queensland Museum 45(2): 266. 2000:- Saxicoline 
members of the Egernia siriolata complex in easlern Australia 
have had a complex laxonomic hislor\\ Cogger (1960) 
demonstrated ecological and morphological separation ol' 
sympatric saxicoline and arboreal species of this complex in 
the Warrumbungle Ranges, NSW, identifying the arboreal 
species as siriolata aw^ describing the saxicoline species as 
E. suxaiilis saxati/i.s. Other saxicoline populations from S!^ 
Australia were described as E. saxafilis intermeiHa, the 
subspecies name referring to the inlomiediale morphology ol' 
these populations, which were allopatric to E striolata. 
Subsequently, some workers have assumed that all saxicoline 
members of the complex in SH Australia, including the New 
Lniiland Tableland, are ,var£///7/.v (see 1 lorton. 1 972; Cogger, 
1988). Wells & Wellington (1984) named an additional 
species, E. mcpheei, from 3 specimens from the Coffs 
Harbour area, NSW, but did not differentiate it from either 
subspecies of E. saxatilis, or resolve the limits of tJie 
distribution of either species. The description of saxatilis 
intermedia refers to two at>pical northern NSW specimens 
subsequently reidenlificd as E. mcphcei Shea & Sadlier. 
1999 for the reideniitlcation and t> pe status). 

A single Queensland Museum record of £". saxatilis was 
reported from SH Qld, close to the NSW border, by 
Covacevich & Couper (1991). The specimen on which this 
record was based was exchanged with the United Slates 
National Museum (IJSNM) in 1 976. and was not re-examined 
prior to publication of the record. 

We here report re-cxaminatioii of this specimen (formerly 
QM J28654, now USNM 2039531, another .specimen from 
nearby, and an unambiguous photographic record from a third 
Queensland iocalilv. and identily all three as Egernia mcpheei. 
■f bus, this species is added to, and £. saxatilis removed from, 
the list ofQueensland reptiles (Covacevich & Couper, 1991 ). 

USNM 203953, preserved 1 Aug. 1 973, is one of a litter of 
more than three bom to a wild-caught female from near 
Girraween, SE Qld, collected 2 Dec. 1972. It is brown dor- 
sally, with 28 midbody scales. 23 lamellae below the foiuth 
toe. and dorsal scales smooth. The snoul-vent length is 
6 1. 6mm, and tail length 79mm (TL/SVL 128.2%). 

On 27 Jan. 1999, imi large adull individuals were closely 
observed on a large rock outcrop at the summit of the 
McPherson Range al Moss Garden (28*' 1 7'S 1 52''26T^) on the 
NSW/Qld border by GS and RS. One (Australian Museum 
R 1 53859) was on the NSW side of the border fence, while the 
other, not collected, was observed one metre distant on the 
Qld side of the fence. The rock outcrop created an exposed 
sunny area in what was otherwise wet sclerophyll/ rainforest. 
The collected specimen has 30 midbody scales, 23/22 
lamellae below the fouitli toe. dorsal scales biuntK keeled, 
snoul-venl length 134mm. and tail length 157mm (TL SVL 
126.6%). 

At least three individuals were observed, and one photo* 
graphed, by RJ on 6 Aprill997, on the southern peak of Ml 
Mitchell, SEQId(28"04\S 1 52^23 T'). inhabiting closed rocky 
grassland with shrubs and grasslrees. above a mosaic of wet 
sclcrophyll/rainforesl. 'fhe photographed indi\idual displa\ed 
dark brown dorsum, brigin orange bod\ venter, and blunth 
keeled dorsal scales. 

Based on unpublished studies by the senior author, E. 
mcphcei is distributed along the coast and E side of the Great 
Dividing Range of NE NS W, from the Bamnglon Tops area to 
the records reported herein. These records are the most inland 
localities known for the species, aiid probably represent a 
migration along the Clarence Rixer valic> from more coastal 
populations. The species is differentiated from the 



e geographically proximate E. striolata in its larger size (SVL 
up to 143mm vs 119mm), more chocolate-brown dorsal 
coloration (vs grey), more brightly coloured \enter (bright 
orange to orange-yellow vs dull orange-bellow to vellow); 
longer tail (tail length = 0.907(snoul-vent'lenglh)''"^^ vs tail 
length = I.352(snoui-venl length)"'"'^'; tail Ienuthas%ofSVL 
111.6-143.0%, mean = 127.1% vs 95.7%-123.3%, mean = 
1 09.6%) and greater number of lamellae below the fourth toe 
(21-28, mean = 24.7 vs 16-26, mean - 20.2). It may be 
differentiated from E. saxatilis by having dorsal scales 
blunll) keeled in adults and smooth in juveniles (as in E. 
striolata-. vs sharply keeled in adults, more weakly keeled in 
juveniles), fewer midbody scale rows (27-32 vs 35-41, mean 
^ 37.0 for E, v. saxa/ih\s\ 30-42, mean ^ 34.5 for E, s. 
intermedia) and a reduced dark upper lateral zone (vs strong). 
1 be nearest populations of E. saxatilis are in the 
Warrumbungle Mlns (£. .v. saxaiilis) and on the Newnes 
Plateau W of S\ dne\ {E, s. intermedia). Egernia mepheei and 
E. striolata, which are allopalrically distributed, both inhabit 
crevices in trees and rocks when in isolation from other 
members of the complex, so that the ecological separation 
observed by Cogger (I960) between E. striolata and E. 
saxatilis in sympatr)' is not applicable in this instance. 

Comparative Material 

/■:. mcphcei: AM 4X73. RSIOS. RII8S'>. R12740. R](t9'Jfl. RI6W2. Rl^'W-*-)?. 
R41 174. R4I815. R418:(). R54308. R54456. R547y7-W. R54807. R59315. R()()4K7. 
Rh2338, R66154-S7. R6823*). R68474-75. R7I400. R76514. R906O2. R93468. 
R%83U. K%«34. R968y4. K977()4. RI08766. RU 1944-46. RI 12279. RI2(H90; 
Northern Tcmior\ Vlasciini R4S08-I(1, 

/:: Mridala: AM RU)5n, RI()54-55. R1499. R1825. R2005. R^8%-98. R4171. 
R9315. R94034)5. RII055. Rlif)9f)a-b. RI1597a-b. RI3899. Ri4%l-67. R152.':0. 
R 1 5254-6(J.Rl 5284-86. R15288. R15290. R15376. R15538-(8. RI555(KS9. R16778-79. 
R17()45-9ft. RI7664-66. R178f«^>-70. R18773. R!89()9-I8. R18924. R2()28l. R203I5-17. 
R2067I. R2073l-3.'5. ri2l448. [i27980-81. R28027-30. R.3n328. R3i.S97-98. R3177n-72. 
R4I801.09. K4I8I1-I4. R4I8I8-24. R4I827-29. R4I831. R4I840-4I. R43439-57, 
R43462-67. R446f)i. R44762. R47338-19. R.'527I7. R52947-50. R5245]. R5776^)-70. 
R57873. R58259. R6(W8!. R6m83-8G. RwWt)4. R604%. R6(>144-t3. R66i48^9. 
R67'>2l-22. R683n. R695S9-tX), R695'>9. R92464. R924^i>-/»(i, R92468. R9247n.74. 
R94534-36. R94727-28. R94783. R9655(l. R9W.28. Ri 1074^1. RI 10755-56. RI 12852. 
R] 12^)53, RI 13322. R]21037^2: AtislniJian Ncjlional Wildlitl- Colleclion R320I. 
R39Cvt-(>7, R4052-55. R4544-56: Mascimi of Victoria D9273-75. 09276-77. D9278-79. 
015423-27. D.54283: QM J5i. J26.3. J4I2-13. J415-22. .11(H87. .113354. .113356-57. 
J13752-70. .114246. J28526-27. J30095-96. .13tl7l7. J30664. .131862. .134125. J34797, 
J354I3; South Australian MiLseum R154I8. 

Acknowledgements 

We ihank Ci. Zug for providing data on USNM 203953. 
Lilerature Cited 

r(X"iCiER. H.G. 1^60. The ecology, morphology, distribution and 
speciation of a new species and subspecies uf the genus 
EgerniiJ (Laccrtilia: SeincidaeK Records of ihe .Australian 
Museum 25(5): 95-105. 
I'>8S. Reptiles and amphibians of Australia. 5th Edn. Pp. xxi, 688. 
(Reed Books: Frenchs Forest). 

COVACEVICH. J. A, & COUPER. P.J. 1991. The reptile records. Pp. 
45-140. In Ingram, G..I. & Raven, R.J. (eds) An atlas of 
Queeiisiimd's frogs, reptiles, birds and mammals. (Queensland 
Museum: Brisbane). 

NORTON, D R. 1972, Evolution in ihe genus Egfrniu (Lacertilia: 
Scincidae). Journal of Hei-pelology 6(2): 101-109. 

SHEA, G.M. & SADEIER. R.A. 1999. A catalogue of the non-fossil 
amphibian and reptile type specimens in the collection of the 
Australian Museum: types currently, previously and 
purportedls present, Technical Reports of the Australian 
Museum 15: 1-0 1. 

WELLS. R. W. & WELLINGTON, C.R. ! 984. A synopsis of the class 
Reptilia in Australia. AusiralianJoumalofl lcrpetology 1(3-4): 
73-129. 

G.M. Shea, Dept of Veterinary Anatomy & Palholo^ry, Sydney 
University, Sydney 2006; R. Sadlier. Australian Museum. 
College St. Sydney 2000; R, Johnson. Environmental 
Protection Agency. PO Box Roma 4455, Australia; 14 
Eebruary 2000. 



BOUNTIANA, A'NEW GENUS FOR ERIPHIA NORFOLCENSIS GRANT & 
MCCULLOCH, 1907 (CRUSTACEA: BRACHYURA: ERIPHIIDAE) 



PETER J.R DAVIE AND PETER K.L. NG 

Davie, PJ.F & Ng, P.K.L. 2000 06 30: Bountiana, a new genus for Eriphla fwrfolcensis Grant 
& McCuHoch, 1907 (Crustacea: Brachyura: EriphiidaeA Memoirs of the Queensland 
Museum 45(2): 267-272. Brisbane. ISSN 0079-8835. 

Eriphia norfolcensis is separated from Eriphia sensu stricto and placed into a new genus 
Bounfiana. The two genera differ in many features including carapace shape, eye size and 
shape; differentiation of etTerent branchial openings; length of second antennular segment; 
length and disposition of basal antennal segment; conformation of gonopod 2; and 
distinctness of suture between male thoracic stemites 2 and 3. A lectotype is designated for E. 
norfolcensis Grant & McCulloch, 1 907. □ Brachyura, Eriphiidae, Eriphia, Norfolk Island. 
Lord Howe Island, Australia, new genus. 

PJ.F Davie, Queensland Museum. PO Box 3300. South Brisbane 4101, Australia: P.K.L. 
Ng. Department of Biological Sciences, National University of Singapore, 10 Kent Ridge 
Crescent. Singapore 1 19260, Republic of Singapore; 21 March 2000. 



Eriphia Latreille, 1817 (type species Cancer 
spinifrons Herbsl, 1785, a junior synonym of 
Cancer verrucosus Forskal, 1775) (Eriphiidae 
sensu Ng, 1998) currently contains seven 
species, viz. E. verrucosa (Forskal, 1775), E. 
gonagra (Fabricius, 1781), E. sebana (Shaw & 
Nodder, 1803), E. smithii MacLeay, 1838, E. 
scabricida Dana, 1852, E. squamata Stimpson, 
1 860, E. granulosa A. Milne Edwards, 1 880, and 
E. norfolcensis Grant & McCulloch, 1907. 
Eriphia verrucosa is found in the Mediterranean, 
E. gonagra is known from the western Atlantic, 
and E. squamata is known only from the Pacific 
coast of tropical America. Three species, Eriphia 
sebana^ E. smithii and E. scahricula al 1 have wide 
Indo-West Pacific distributions. The remaining 
species, Eriphia norfolcensis is only known from 
Norfolk Island, Lord Howe Island and the coast 
of New South Wales. Examination of a series of 
specimens of E. norfolcensis shows that this 
species is aberrant within Eriphia, and more 
closely allied to Globopilumnus Balss, 1933. A 
new genus is here established for E. norfolcensis. 

Measurements provided are of the carapace 
width and length respectively (including spines). 
G 1 and G2 are abbreviations for male first and 
second gonopods respectively. Specimens are 
deposited in the Australian Museum (AM), Syd- 
ney, and Queensland Museum (QM), Brisbane. 

ERIPHIIDAE Alcock, 1898 
Bountiana gen. nov. 

TYPE SPECIES. Eriphia norfolcensis Grant & 
McCulloch, 1907, by monotypy. 



ETYMOLOGY. Named after H.M.S. Bounty, in 
remembrance of the group of mutineers who lived 
undiscovered in the South Seas for so long. The ancestors 
of the mutineers were, many years later, moved from 
Pitcaim Island to Norfolk Island where their descendents 
live to this day. 

DIAGNOSIS. Carapace transversely ova!, 
strongly vaulted anteriorly; regions poorly de- 
fined. Front obliquely detlexed, with transverse 
ridge either side of notch. Anterolateral margin 
with three small, blunt, well separated, spines. 
Eyes relatively small with small corneas (see Fig. 
3A). Efferent branchial openings not distinct, 
almost completely covered by third maxillipeds, 
endostomial ridges weak, not forming circular 
opening with epistome; antero- external margin 
of third maxillipeds rounded; third maxillipeds 
gaping. Second antennular segment very short, 
less than half length of basal segment. Basal 
antennal segment with outer distal lobule just 
touching front, and placed close to inner angle of 
orbit (Fig. 3A); orbit closed. Chelipeds 
asymmetrical. Fingers of chelipeds gaping. 
Ambulatory legs short, stout; dactyli very short. 
Gonopod 2 flagellum subequal in length to basal 
portion. Suture between male thoracic sternites 2 
and 3 distinct. 

REMARKS. The ovate, longitudinally strongly 
convex and bulging carapace of Eriphia 
norfolcensis immediately separates it from 
typical Eriphia species, which are hexagonal or 
transversely hexagonal, and only moderately 
convex. This alone is sufficient to necessitate the 
establishment of a new genus, Boimfiana, for 
Eriphia norfolcensis. There are also, however. 



268 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. \ , Bountiana norfolcensis (Grant & McCulloch, 1907), lectotype male, 12.2 x 18.1mm, Norfolk Island; A. 
dorsal view; B, ventral view. 



many other characters of generic significance and 
these are summarised in Table 1. All species of 
Eriphia, including the type species, E. verrucosa 
(Forskal, 1775), have been examined and taken 
into account when constructing Table 1; this 
material forms part of a separate revision of 
Eriphiahy S.K.Koh& P.K.L. Ng(unpubl. data). 



Botmtiana norfolcensis is closer in general 
appearance to species of Globopilumniis, the 
only other genus within the Eriphiidae. It can be 
effectively separated from this genus because in 
Globopilumnifs the supra- and infra-orbital 
angles do not meet, such that the antenna enters 
the orbit through an orbital hiatus. 



A NEW CRAB GENUS, BOUNTJANA 



269 




FIG 2. Bountianu norjolcemis (Grant & McCulloch, 1907), femate, 16.5x1 1.8mm, Norfolk 
Island; A, dorsal view; fl, firontal view showing claws. 




FIG 3. Orbil and frontal regions; A, Bomtiam norfnlcensis (Gram & McCulloch. 1907), male, 1 6.3 x^l 1 .2mm, 
QMW2490?;B,£/-^/iWiycfli?r/£?«/oDana, lS52,male, i9.3x 13.3mm, QMW 1211 7,Lad> Ellioilsland, SliQld. 
(b.a.s. ^ basal antennal segment; j.O. - imief orbital angle). 



270 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4. Third maxillipeds; A. Bountiana norfolcensis (Liiant & McCulloch, 
1907), female. 1 6.5 x 1 1 .8mm. QMW24962; B, Eriphia scahricula Dana, 
1852, male, 19.3 x 13.3mm. QMW121 17, Lady Elliot Island, SE Qld. 



Bountiana norfolcensis 

(Grant & McCulloch, 1907) 
(Figs K2,3A, 4A,5) 

Eriphia norfoicen.si.s Grant & McCulloch, 1907; 151. pi. 1: 
McNeil & Ward 1930: 38 i; Holthuis, 1968: 218. 

Pseiidoziiis sp. Bennett, 1964: 67-68, tigs 62-68, 132; Dell, 
1968: 17-18; Guinol, 1968: 330-331. 

MATERIAL. LECTOTYPE: AMG5827, 6 (12.2 x 
18.1mm), Norfolk I., Tasman Sea, 29°02.5'S 167°57'E, 
EE. Grant. PARALECTOTYPES: AMG5827, 9 
( 1 0.8 X 15.7mm), 1 juvenile (7.5 x 10.0mm), Norfolk I., 
Tasman Sea, 29*'02.5'S 167"57'E, RE. Grant OTHER 
MATERIAL: AMP446, ?, AMP448, 9, AMP449, 6. 
Lord Howe I., no other data. AMP4032, S 
(7.6X 11.0mm), 9 (9.8 x 14.4mm), AMP5254, S 
( 1 0.0 x 14.6mm), 29(10.7x15.6, 8.7x 12.6mm), Lord 
Howe L, 3P33'S 159WE, E.A. Briggs. AMP5255, 



2 9 , AMP5256, 5 <5 , 9 , Lord Howe 
L, A.R. McCunoch. AMP10328, J, 
9, Lord Howe I., reef, April, 1932, 
A.A. Livingstone. AMP6285, 
AMP6842, ovig. 9,AMP7889, 9, 
Shell Harbour, NSW, 34°35' S, 
150°53T, 1923, G. McAndrew. 
AMP 11285, (5, Harbord, NSW, 
33°47' S. l5ri7^E, 24.11.1947, E. 
Pope.AMP17289,29 (18.9x 13.0; 
10. 9x7. 8mm), 3<? (9.3 x6. 7; 
11.5x8.3; 18. 2x 12.4mm), 
Slaughter Bay, Norfolk I., imder 
coral aibble and weed, outer reef 
crest, low tide, 21.03.1969, D.J. 
Griffin. QMW24902, 2 9 
(16.5x11.8; 18.5X 12.8mm), 2S 
(! 6.3 X 1 1 .2; 1 8.3 x 12.6mm), same 
data as AMP 17289. 



DESCRIPTION. Carapace 
transversely ovate, strongly 
vaulted anteriorly; dorsal 
surface smooth except for 
scattering of low tubercules anterolaterally; 
regions poorly defined, gastric region weakly 
separated by grooves. Front with transverse 
ridge, de flexed downwards, slightly 
denticulated. Eyes relatively small with small 
corneas. Infraorbital margin with about eight 
denticles. Anterolateral margin armed with four 
widely separated, low tubercles. Second 
antennular segment very short, less than half 
length of basal segment. Antennae very short, not 
reaching orbital margin; basal antennal segment 
with outer distal lobule just touching front, and 
placed very close to inner angle of orbit. Third 
maxillipeds gaping medially; outer surfaces 
relatively smooth, pubescent with long stiff 
setae; merus irregularly pentagonal, slightly 



TABLE 1. Differences between Bountiana gen. nov. and Eriphia Latreille, 1817. 





Bountiana 


Eriphia 


Carapace 


strongly vaulted anteriorly 


moderately convex to almost flat anteriorly 


Eyes 


relatively small with small corneas (see Fig. 3A) 


large, corneas bulbous (see Fig. 3B) 


Carapace shape 


transversely oval 


sub-hexagonal 


Efferent branchial openings 


not distinct, almost completely covered by third 
maxillipeds (Fig. 3 A), endostomial ridges weak, not 
forming circular opening with epislomc 


very distinct, not covered by third maxillipeds (Fig. 
3B), endostomial ridges strong, forming circular 
opening with epistome 


TTitrd maxilHpeds 


antero-extemal margin rounded (Fig. 4A) 


antero-extemal margin sub-auriculifomi (Fig. 4B) 


Antennules 


second segment very short, less than half length of 
basal segment (Fig. 3A) 


second segment long, three-quarters or more length 
of basal segment (Fig. 33) 


Antennal position 


basal antennal segment with outer distal lobule just 
touching front, and placed very close to inner aiigle 
of orbit (Fig. 3A) 


basal antennal segment with outer distal lobule not 
touching front, and widely separated from inner an- 
gle of orbit (Fig. 3B) 


Ambulatory dactv lus 


short and stout 


long and slender 


Gonopod 2 


flagellum subequal in length to basal portion 


flagellum distinctly shorter than basal portion 


Male thoracic stemites 


suture between stemites 2 & 3 distinct 


suture between stemites 2 & 3 not discernible 



A NEW CRAB GENUS, BOUNTIANA 



in 




FIG. 5. Bountiana norfolcensis (Grant & McCulloch, 
1907), lectotype male, 12.2 x 18.1mm, AMG5827, 
Norfolk Island; A, enlargement of apex of gonopod 1 ; 
B, gonopod 1; C, gonopod 2. 



notched at distal outer margin, antero-extemal 
margin roimded, not sub-auriculiform; with a few 
long, stiff setae. Efferent branchial openings not 
distinct, almost completely covered by third 
maxillipeds, endostomial ridges weak, not 
forming circular opening with epistome. 
Pterygostomial region smooth. Suborbital region 
relatively smooth. 

Chelipeds markedly dimorphic; inner margins 
of all segmeiits pubescent; distal end of merus 
pubescent, surfaces smooth. Distal blunt spine 
present at ventral surface of basis-ischium. 
Carpus with acute spine on median- inner margin, 
smaller one present ventrally; surfaces 
tuberculated anteriorly. Chelae relatively short, 
stout, with longitudinal rows of tubercles. 
Pubescence present on entire dorsal surface of 
palm, less dense on ventral surface. Small basal 
non-molariform tooth present on dactylus. 
Fingers of major chela strongly curved, closing 
with a prominent gape. Cutting edges of lingers 
of minor chela minutely denticulated. Female 
claws similar to male. 

Ambulator)' legs short, stout, smooth, both 
anterior and posterior margins with prominent 
long and short setae; merus with minute tubercles 
anteriorly. 

Anterior male thoracic sternites smooth. 
Abdominal surfaces smooth. 01 short, stout, 



broad at base tapering distally, tenninating in 
broad tip; long stout spinules present on distal 
half of inner edge, longest medially; minute 
spinules on outer surfaces over distal third. G2 
relatively long, slender; distal half forming 
curled flagelium. 

REMARKS. Since Grant & McCulloch (1907) 
described the species from Norfolk Islands, there 
have been only two subsequent reports bv 
McNeil & Ward (1930) and Holthuis (1968). 
McNeil & Ward (1930) added Lord Howe Island 
and New South Wales as localities for this species. 

The Pseudozius sp. of Bennett (1964) from 
Campbell Island, south of New Zealand, is 
without doubt synonymous with Bountiana 
norfolcensis. This was tlrst noted by Guinot 
(1968: 330) whilst reviewing Pseudozius and we 
concur with her conclusion. Furthennore, Dell 
(1968: 17) provided strong evidence that this 
species could not have come from Campbell 
Island, and must have been collected from an 
unknown locality elsewhere in the Pacific. This is 
based on the fact that it has not been recorded 
since, despite subsequent extensive collecting 
expeditions to the island. Also it w as apparently 
collected during a trip by the government vessel 
Hinemoa whose captain, Mr J. A. Bollons, was 
notoriously inaccurate in recording where 
specimens were found. 

Grant & McCuUoch's (1907) original 
specimens (AMG5827) were examined and a 
lectotype male (12.2 x 1 8. 1mm) is here designated. 

DISTRIBUTION. Norfolk Island, Lord Howe 
Island, and New South Wales, Australia. 

ACKNOWLEDGEMENTS 

We thank Penny Berents and Roger Spring- 
thorpe of the Australian Museum, Sydney, for 
assistance during our respective visits, and loan 
of specimens. Also we express our gratitude to 
S.K. Koh and S.H. Tan for help with photographs 
and drawings. 

LITERATURE CITED 

BALSS, H. 1933a. Beitragc zur Kenntnis der Gattung 
Piliimnns (Crustacea Dekapoda) und vewandter 
GattLingen. Capita Zooloeica 4(3): 1-47, 

BENNETT, E.W. 1964. The Marine Fauna of New 
Zealand: Crustacea Brachyura. New Zealand 
Oceanographic Institute Memoir No. 22. Bulletin. 
New Zealand Department of Scientific and 
Industrial Research 153: 1-120. 

DANA, J.D. 1852. Crustacea. United States Exploring 
Expedition during the years 1838, 1839, 1840, 
1841, 1842 under the command of Charles 



272 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Wilkes, U.S.N. Vol. 13: 1-685 (1852), 686-1618 

(1853), Atlas (1855): 1-27. 
DELL, R.K. 1968. Notes on New Zealand cfabs. 

Records of tbe Dominion Museum 6(3); !3-28^ 
FABRICIUS J,C. 178L Species Insectonjmodnben^ 

eorum Differontias, spectficas,. SyiKimyma 

auctorum, iMk hat^i^ Metamoiphosia adectis 

Observationibus, Descriptionibus, Vol. 2. 

(Hambuig & KJlonii). 
FORSKAL, p. 1775. Descripliones animalium avium, 

amphibiorum, piscium. insectorum, veimium; 

quae in itinere oriemali observavitPetru&FoEskEU. 

(Hatniae). 1-19. 1-XXXlV. 
GRA^T. F.n. & MCCULLOCH. A.R. 1907. Decapod 

Crustacea from Norfolk island. Proceedings ol lbe 

Linncan Socie^ of New South Wales 32(1); 

151-6. 

GUI'NOT, D. l%Sa. Recherches preliminaires sur les 
groupenienis nalurels chez les Crustaces 
Decapodes Brach} cures. VI. Les Carpilinae. 
Bulletin du Museum National d*Histoire 
Naturelle, Paris Ser. 2, 40(2): 320-334, 

HOLPHUIS. L.B. 1968. Are there poisonous crabs? 
Crustaceana 15(2): 215-222. 

LATREILLE, P. A., 1817. Nouveau dictionnaire 
d^histoire naturelle, appliqude aux arts, a 
ragticultt^ k r^conotnie rurale et domestique, k 
!a m^decifie» etc, VoL W. 

MACLEAY, W.S. 1838. On the Brach>airous Decapod 
Oustaceabrought from the Cape by DrSmith. Pp. 



63-72. In Illustrations of the Annulosa of South 
Africa; being a portion of the objects of natural 
history' collected during an e.vpedition into the 
interior of South Africa, under the direction of Dr 
.AAdfew Smith, itilheyears 1834, 183$, and 1836; 
fitted out by 'The Cape of GoodHobe Assc^iatioa 
for Exploring Central Africa*. Illustr. Zool. S. 
Africa Invest. (London). 

McNElLL, F.A. & WARD, M. 1930. Carcinological 
Notes. No.l. Records of the Australian Museum 

17(9): 357-83. 

NG, R1C.L. 1998. Crabs. Pp. 1045-1155. In Carpenter, 
K.E. & Niem, V.H. (eds) FAO Species 
identification guide for tishery purposes. The 
.living marine ix^durces of the Westero C^tr^l 
F^inc. \bliime 2^ C>phalopod5, ci*U5laceansj 
holothuna&s and shades, ^ood and Agriculture 
Orgariisation: Rom6). 

SHAW, a & NODDER, P.P. 1803. Vivarium Naturae, 
sive rerum naturalium. variae el v ividae icones. ad 
ipsam naturani depictae el descriptae. Crustacea. 
Pis 589-61 2, unnutnbercd pages. In The 
naturalist's miscellany, or coloured ilgures of 
natural objects, drawn and described immediately 
from nature. (London). 

STIMPSON, W. 186Q. Notes on Nordi American 
Cr^sta^a, in ihe Mu$eutn of the Sndthsonis^ 
Ir^titiition. No. 11. Annuals of ths Lyceum of 
Natui^History ofNew York?: 177-246 (49-1 18). 



MALARINA. A NbW SPIDER GIN US (ARANHAH: AM \L ROBIOIDEA; 
KABABININAE) FROM TTIE WET TROPICS OF QUEENSLAND, AUSTRALIA 



VALEWE tODD JOAVtBS AND fe^jRlSIINE L. LAMBKIN 

DjAies, \Vr. ft Lambkin. C.L. 2000 0(> 30: Mularinu. a new spider genus (Araneae: 
Amaufobioidea; Kababinii^aej from the Wei Iropics ot QueeasJand, Ausiraiia. Memoin oj 
the;Q^starurmsevm 45(2): 27M853risbimejlSSN 0079^8835. 

Four dpcCteSpfMjfiJW'Wii gen. nov. tu-e described indicating the lot ui endemisni of species in 
North Queensland. These are M monteith'ty hft massey^emis, M coUina and M- canhvcll, \ 
cladisiic analysis suggesth ihai i)ie Kabafcariinae fbmia weli supported monophyleiic grcnip^ 
tt^)Hg^ it^placeinentinafamily remainsprpblematical.OAfo/wvVi^, Aranme. i<^bobmimB, 

Vakrie TodttDavies, Qus^nstond Museum, PO Box SMI, South Brisbane 4 101: Chrfsdne 
Xj L0airkfrK P^a^pmttwiQdiosy^ ^ntomahe^^ University ofQueen&ltmL St Lucia 
402:Ausfrath: ifMarcH. 1999. 



Maliirifia is the third genus to he described in 
the K-ihahininae. ihc LUhers being Kuhahina 
Davies, 1995 j^nd Cc^rbm^ Oavies. 1999. For 
nomendatiu'aliwposes-Omti^^^ the 
aothorof 11h]^l!iew ge^us and its species' names. 

METHODS 

All the spideis am from ramiprcste in the Wet 
Topics region ofNoirSi Que^fislMd f)etwMn 

latitudes P"16'and 1S°36'S. Collection methods 
include liller-sieving followed bv heal e\traction 
\v\ funnels, pit-fall colleetirm, pyretiiruin spraying 
of tree-trunks and fallen logs, hand collecting 
from under logs in daylight and night collecting. 
Co-ordinates arc given in square brackets when 
these are not given in the original data. 
Measurements are in niiHimcires. Notation of 
spines follows Platnick <fc Shadab (1975). The 
illustrations were drawn with the aid of a camera 
Uieida; the Jjeft inale jpalp is MlMS^ted. All njaterial 
is lodged fit thtf Qt£eti»lMd MtiSdim f QM^^ 

Table I lists anattDOUCil abbreviations used in 
the text in Fabte 3; abbreviations on ill^s- 
trardotiis are plained in the legends to figures* 
Collectors: DC, D. Cook; DY D. Yeates: GBM, 
G.B. Monteith; GT, G. Thompson; HJ, H. 
Janetzki; RR, R. Raven; .5H> Hatnlet; VEfJ, 
VE. Davies. 

SYSTEMATTCS 

KABAMINTNAF 

DJ^GNOSTS. Epigynum with medial atrium 
CjJfeviously rcferrcdto as *fossa'), which is wider 
than long; spermathecae posterior fir lateral to 
atrium. Male palp with rounded legulum with 
prolateral groove; ihe course of sperm 



showing clearly. Membraneous conduelor: 
medial! apophysis absenl. Tibial apophysis with 
ventral anddorso-retroiateral branches. Posterior 
spinnerets ioing ^^th. lender teftnitial iie.ginciit 

(Fig. 

DESCRJPTION. Three clawed. C^W^flpWe 
highest in foveal region (Fig. IB); po$feifi5r 6yfc 

row straight or slightly reeur\'ed; AME reduced 
("Fig- IC')- Chelicera with two retroniargmal and 
n\o proniarguia! teelh (fig. 1!.): ptoiaterul 
filamentous seta at base of fang longer than other 
setae. Labium about as wide as long: sternum 
pomfed posteriorly (Fig. ID). Legs 1423; 
ieaiher} hairs, ridged cuticle. Tarsal tnchobolhria 
in a smgle line increasing in length disially; 
bothnum collanl'oim. Tarsal organ slit-like 
broadening distallv- Male palpal embolus with or 
without trcixifoal embolic appptorsis-Ciibe^lupi 
(two fields) present oraTjscjit iir fetngli^srabs^ 
ill males; proximal calamistrum N\*ith one row of 
setae: large broad colulus present \\ hen cribcllum 
is absenl. Two major ainpuUatc gland spigots of 
unequal ^ize on female ALS, one and a nubbin in 

Malarfna geti. tiov. 

ETYMOLOCN; Derived Irom ifit Alvariginal word, 
THolar^ iUeMng solder's web in the Dyirbal tangu^ of 

DIAGNOSIS. Cribellale spider (cf Carhim'u) 
with paracribeliar spigots on female PMS. 
Epigynum ha\ uig a posterior knob and nantuv 
postero-lateral insemination ducts (cf. Kababbia 
which lacks the well-marked epigynaJ knob and 



274 



MhMOiRS OF Tilt: QULENSLAND MUSEUM 





FIG 1. ^ilaiarimm(>nteithi9pyloy' A-I, 9 ; A-C,carapaec (dorsal* lateral^ frontal); D, enditesandstertium; 
E, chelicem; F.flpjnn^r^ {lateral); G-I, epigynum'{vetitwl, dorsal, l^rai). J-L. cJ ; J. palp (ventral); K, tibial 
apophysis (retrolateraU docrsal). 

and conductor arising antero-ventrally on loi^ ventral selae reacliing about half way up 
tegulum (Fig. 5A); the embolus having an Qnnbiiim (Fig. IJ) aad a small sub-cental 
elaborate embolic apophysis like Carbinea 6ut prolatero-dotsal spine (Fig. IL); the: tibial 
here it is unbranched. Male palpal tibia with 2-4 apophysis is dorso-r&tiX)Iateral. 



NEW SPIDER GENUS FROM THE WET TROPICS 



275 



TABLE 1. List of anatomical abbreviations. 



A r 


abdomen length 


ALE 


anterior lateral eyes 


ALb 


anterior lateral spinnerets 


AME 


anterior median eyes 


APOPH 


apophysis 


AW 


abdomen width 




calamistiiim 


CB 


cymbium 


CH 


cheliceral 


r^T 


carapace length 


CK 


cnbellum 


L^Vv 


carapace width 


b 


embolic 


brio 


epig\'nal 


ID 


insemination duct 


MAP 


major ampullate spigots 


MT 


tnptntnrsfll 

11 IVlClLCU JCll 


PGR 


paracribellar spigots 


PLD 


prolaterodorsal 


?IE 


posterior lateral eyes 


PLS 


posterior lateral spinnerets 


PME 


posterior median eyes 


PMS 


posterior median spinnerets 


RTA 


retrolateral tibial apophysis 


T 


tarsal 


TRICH 


trichobothria 



Malarina monteithi sp. nov. 
(Figs lA-L, 2A,B,6;Table 2) 

ETYMOLOGY. For Dr GB. Monteith, entomologist, who 
has collected widely in the tropics of N Queensland. 

MATERIAL. HOLOTYPE: 9, Malanda Falls, I7°2rS, 
145°35'E, N Qld, 750m, pyrethrum spray logs and trees, 
31 Dec. 1989, GBM(QMS35253).PARATYPES:N Qld, 
2 c?, same data as holotype (S35254); 9, 2 c5, Mlllaa 
Millaa Falls, 17°28'S, 145°36^E, 800m, sieved litter, 17 
May 1995, GBM (S35255); 9, The Crater Nat. Park, 
17'^26^S, ]45°29'E, 950m, pyrethrum logs, 28 Dec. 1989, 
GBM (S 35256); <5, 3 9, Maalaji State Forest (17°35'S, 
145°35'E) in and under logs, 20-24 April 1978, VED. RR 

(535257) ; 9, Majors Mtn (17°38'S, 145°32'E) same data 

(535258) ; 9, Mt Fatlier Clancy, Maalan, litter, 21 April 
1978, RR(S35259); 9, Ravenshoe, 17°39^S, 145°30'E, 
920m, pitfall, 1 Dec.1997-5 Feb. 1998, GBM, DC (S39202); 
d. Red Rd tumoff. 17°49'S, I45°33T, TuUv Falls Rd, 
pitfall, 8 Dec. 1989-5 Jan. 1990, GBM. GT, HJ (S35263). 

DIAGNOSIS. Small (2.8-3.6) cribellate spiders; 
proximal calamistrum. The insemination ducts 
are simple. The embolic apophysis has two long 
setae extending beyond the rest (Fig. 2B). 

DESCRIPTION. Female. CL 1.4, CW 1.1, AL 
1 .7, AW 1 .2. Carapace with two dark longitudinal 
bands (Fig. I A); highest at fovea. Viewed from 
top, eye rows straight. Ratio of AME:ALE: 
PME:PLE is 6:10:10:10. Legs 1423 (Table 2), 



TABLE 2. Palp and leg measurements (mm) of 
9 ( 6 ) Malarina monteithi sp. nov. 





Femur 


PatelW 
Tibia 


Metatarsus 


Tarsus 


Total 


Palp 


0.6(0.8) 


0.6 (0.9) 




0.5 (0.9) 


1.7(2.6) 


Legl 


, 1.5(1.8) ^ 


2.0(2.3) 


1.4(1.9) 


0.9(1.2) 


5.8 (7.2) 


II 


1.3(1.5)^ 


1.6(1.8) 


1.1 (1.4) 


0.7 (0.9) 


4.7 (5.6) 


HI 


1.1(1.4) 


1.4(1.5) 


1.1 (1.3) 


0.7 (0.7) 


4.3 4.9) 


IV 


1.5(1.8) J 


1.9(2.2) 


1.5(1.9) 


0.8(l.q)j 


5.7 (6.9) 



banded. Notation of spines: Femora, 1, DOlO, 
PO 1 0; 11, D II 0, POO 1 ; III, D 1 00, POO 1 ; IV, Dll 0, 
POOL ROOl. Patellae, I, DOOl; IL DlOO; III, 
DOOl; IV, 001. Tibiae:!, V020; II, VOlO; III, 
DlOO, POOL ROll; IV, DlOO, POl 1, VI 10, ROl 1. 
Metatarsi, all spined with a distal whorl of 4-5. 
Epigynum (FigslG-I) short insemination ducts 
with anterior loop to spermathecae. These are 
large, together exceeding the width of the atrium. 
Cribellum with two fields; ALS with two major 
ampullate spigots and about 20 pirifom spigots 
and some tartipores; PMS with an anterior minor 
ampullate spigot, and about 12 other spigots - 
two cylindrical spigots (mesal and posterior), 
four smaller paracribellar spigots with strobilate 
shafts and about six acinifomi spigots. PLS with 
spigots of two sizes. Females varied in length 
from 2.8-3.6. 

Male, CL 1 .5, CL 1 .2, AL 1.7, AW LL Coloration 
and eyes like female. Legs 1423 (Table 2). 
Notation of spines: Femora, I, DllO, POlO; II, 
DUO, POOL ROOl; III, DllO, POOL ROll; IV, 
DlOO, POOL ROOL Patellae, L DOOl; III, DOOl; 
IV, DOO 1 . Tibiae, L D 1 00, PO 1 0, V020, ROO 1 ; II, 
D00LP011,V020,R001;III,DI01,P011,V1I1, 
ROll; IV, DlOl, poll, VIU, ROIL Metatarsi 
spined, with distal whorl 4-5. Male palp (Fig. 
1 J-L), spemi duct looping over retrolateral tegulum 
and forward again to base of embolus. Embolic 
apophysis fringed with a prolateral cluster of 
hair-like setae and five plate-like setae terminally 
(Fig. 2A,B). Under (strictly dorsal to) these there 
is also a row of straight setae, two of which are 
longer than the rest and protrude beyond them. 

DISTRIBUTION. Collected from sites on the 
Atherton Tableland (Fig. 6). 

Malarina masseyensis sp.nov. 
(Figs 2C,D, 3A-F, 6) 

ETYMOLOGY From the type locality, Massey Range. 

MATERIAL. HOLOT^TE: 9. Massey Range, 17°I6^S, 
14.S"49'E, 1250m, sieved litter, 10 Oct. 1991, GBM, HJ 
(QM S35260). PARAT\TES: ^ , ? , same data as holot>pe 



276 MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 2. A-D, 6 Malarim spp.nov. embolic region of 6 palp. A, B, M monteiihi (ventral, prolateral); C, D, M. 
masseyemis sp.nov. (ventral, prolateral). c ^ conductor, e = embolus, ea = embolic apophysis, k = keel, ps = 
plate-like setae. 



(S391 85); 9 , Massey Range, 4km W of Centre, Bellenden 
Ker, 1250m, 9-11 Oct. 1991, GBM, HJ, DC (S35261); 9, 
Bellenden Ker Ranae Summit, 17°16'S, 145°5rE, 
1 560m, in litter, 28 Oct 1 983, GBM, DY, GT (S35294); 9 , 
Bellenden Ker, Cable Tower 3, 1054m. under logs, 17-24 
Oct. 1981, Earthwatch/QM, (S35295); Bellenden Ker 
Range, pitfall trap, 500m, 17-24 Oct. 1981, 
Eaithwatch/QM (S 39186). 

DIAGNOSIS. Larger spider (9 's 3.9-4.7) than 
M monteithi. Epigyniim with simple insemin- 
ation ducts; spennathecae togelhernot exceeding 
the w idth of atrium (cf. M monteithi). Palpal tibia 
with two large distal prolaterodorsal spines (cf. 
M monteithi) as well as the sub-centrai spine. 

DESCRIPTION. Female. CL 2.1, CW 1.5, AL 
2.3, AW 1 .3. Carapace without dark longitudinal 
bands. Eyes similar to M monteithi. Legs 1423: 1, 
8.6; II, 6.7; III, 6.3; IV, 8. 1 , not markedly banded. 
Epigynum (Fig. 3A-C) with simple insemination 



ducts with a transverse loop before entering 
spennathecae. Length 3.9-4.7. 

Male. CL 1.9, CW 1.2, AL2.0, AW 1.3. Legs: I, 
9.1; II, 7.0; III, 6.3; IV, 8.6. Palp (Fig. 3D-F): 
large conductor; embolic apophysis curves 
strongly with marked keel; plate-like setae absent 
(Fig. 2C,D). Palpal tibia with two stout pro- 
laterodorsal spines distally (Fig. 3F). Length 3.2-3.9. 

DISTRIBUTION. Collected from sites on the 
Bellenden Ker/Massey Range (Fig. 6). 

Malarina collina sp.nov. 
(Figs 4A-D, 5A,B, 6) 

ETYMOLOGY. Latin C0///5, a hill, referring to the location. 

MATERIAL. HOLOTYPE: 9, Palmerston Nat. Park 
(17°34'S, l45°4rE) under logs, July 1992, J. Wunderiich 
(QM S35262). PARArtTES: 26, 29, same data as 
holotype (S39187); c5. Upper Boulder Ck, 8km N. Tully, 
17°50'S, 145°54'E, 250m, pitfalls, 4-7 Dec. 1989, GBM, 



NEW SPIDER GENUS FROiM THE WET TROPICS 



277 




FIG 3. A-F, Malari/ia musse^ensis, A-Q ^igs^um ^mttia^ tJcir^al, lateral); D-F? (J palp; D, E ( venttel^ 
retroiateral); tibial apophysis (dorsal). 



GT, MJ (S35264): Boulder Ck via Tullv. 17"50'S. 
145°54'£, 650ni, sieved litter, 27 Get 1983, GBM, DY. 

DIAGNOSIS. Small spiders ( 9 :.4-3.6). 
Insemination ducts coiled (cf. A/, monteithi, M. 
masseyensis). Tibial apophysis shorter and 
broader than M monielihi (Fig. 4D). All setae oo 
^bt^ apophysis abtmt same!' TengUl (c{l M 



Diunlf'uhf). Without two long prolaterodorsal 
spines on palpal tibia (cf. M. niasscyensis). 

DESCRIPTION. Female. CL J. 5, CW J,a, AL 
1 .8, AW Ul. Less, 1, 6.7; 5.3; m, 4:7; 
Epigynura (Fig* 4A-G) Wifilf coiled insemination 
ducts. Length 2.4-3.6 

Male. CL 1 .5, CW L L AL 1 .4, AW LO* Legs, 1, 
6.5; n,5.>^flr^+.5; IV, 6.4. Palp (Figs 4D,SA,B), 



278 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4 . A-I, Malarina spp. nov. A-D, M. collina sp Jiov.; A-C, $ ^ epigynum (ventral, dpriial, lateral); D, d M tibial 
apoph\ sis. EA,M. Cardwell sp. ijov,; E-H, eptgyi^m (Veiiti^V Yenlral deared^ di)rsalt Wtisid); paJp 

(ventral). 



NEW SPIDER GENUS FROM THE WE P 



279 




FIG 5. A-KMctlaHmspp. A,B, M. coUina ^; A, 6 palp; B, embolic region (ptblMtpral). C-f , M. cardweB;0-E, 
5 spigots; C, ALS O*?]^); D, PMS; E, paracribelfer^pigpts (pc) on PMS. F, d? palp, tibial apophysis. 



280 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE X Characters and character stalie5,^**B4ulli- 

siaie diaracicr treated as unordered. 



1. AAlIi; .la Uge or larger than ALt- (0)r smallci ( 1 ) 

2. Reirnmarginal CM teefli: 2 < 'Oi; 2 i 1 i: 1 \2\. i3) 
PromMrginal CH teeth: 3+ fU); ( 1 ); 2 (3) 

4. Lunuprolalcral seta at bascDt fanii: absent (0);prc$cjlt( 1 1 

5. Large frontal CH scia: absent iO); present ( 1 ) 
(i. CH lamina: absent (0); present ( 1 ) 

7. Koveal area highest: absent (0); prc&cni H) 
& $ leglrshpnerthaiileglV (0): equal imqr lpngBr^tUT leg 
lV(>> 

9. StKdulaliiiyjrUI&e^Oa a ^COtglt gil30(lt^Pl 

present ( I ) 

IJ - I'eaihcn hairs: abscm pivs^-ni i, I I 

12. MT preening comb: absent (0): present ( Ij 

13. MT TKICH: 2+ (0); 1(1) 

14. T rRICll OdO. ?i (I):doub!ermv (2) 

15. X rod: absent (i ii; pre-icnl i 1 1 

Ift. CK tpUMiing lieMi.: 2 tl»V I (Ihabbcnt {2) 

1 7.^ CR !tpigc»ts£ ubseni tO); In^GgjrtadiMbf ribbed i I ); 

annulate 42) 

1$,* CAl4iflbe&M(U)i cmlDnt^tB (2) 

20. MAP V ALS; mflsal anterSFrl^ 

11. PGR 9 PM8^tme^a¥petbaW(0)| morc(!iaQ^CTfl^ 

1 1 ): absent (2) 

22. Li^iG gonopori!.s: abseml (t)); present { I ) 

23. Medial EPIG atrium^ -absent (0); present ( 1 1 

24. [D:iteettl(^};s^fflpj[K(])f((K>sc;lycoilcdj(2);^^^ 

25/^ Posterior hPU . ^eapB^a^)fi^^IJt^■0);^h^«(l1;E^ 

long {}): stnaJl knob (4> 

26. RPKTi aceiliilar lateral projeetiony ; absent i (J j; present ( I ) 

27. E direction :straigb|(0); clockwise (I );anii-clockiA'ise (2) 

28. Proxinial E APd^U: at|aBm(0),''antirancbcd ( 1 ); bmncbed 

2S». U APOPH 2-3 long s?ia^ dbiEJlt Ift); Wed6nl {1 >, 

30. * E APOPH p(atc4ikc setae.- abs<?n( (01; smaU (0 >, large ( 2 ) 

31 . Pl.D setae E AJ WH; absent (0): present CD 

32. Pjil- process: present (0): absent ( I ). 

33. * Conductor: absent (0); roimded ( 1 ); largp T-^ujicd (2); 
s-sh,*iDetl - laleifon-n (3) 

34 2 eonduetor: absent (0); present ( I) 

35. Median AP(»PH: absent (0); present (I) 

36. Orientation ol'CB to huJh: dorsal (01, rnc<^al ( 1 ) 
^7 P.Mrae\nnbiiim: absent (0): present ( ) ) 

3R. r<T/\loL B length: absent fOi;qnarlLi - m li:v,| 1 ), third (2|; 
hall (3). more than hair(4} 

39. RJ A dorsal branch: absent lO). preseul 1 1 ) 

40. R'l A Iftleral edge: sn-arght (0); miuMicd ( I ) 

41. Pa]pai tibia \vith 2 stout dorsal spines; ab.KtfUl(O); parent 

^ fUpflJ pafella) ap6pw: mm {0)kvttAciA (it 



all s^ta^ Qj^ $m^l2< ^Qphy^isi about same 
length. Sfoad WtlflUs. AlS Wfh 6m major 
ampullate ^igfSt and nubbin, about 17 piriform 
spigots ana some tartipores. FMS with large 
anterior spigot (minor ampullate)«dd I6spjfe)5s 
of uneven size. Letigth 2.6-3.1 

DISTRIBUTION. Collecied ut lower altitudes at 
Palmerson Nal. Park and tiic Walter Hill RMg? 
(.Kg, 6), 



145 





UMalarina montei^ 
MMatarihB ^rd&eli 



147 «, 



riCi. 6. Map of North Queensland sthowing 

distribiitl^ OfA^lmm spp, 



Malariaa cardwetl sp,nov 

ETYMOI-OG^'. Por the locality; C;ud\\ ell Rimge. 

MATERIAL. HOLOTYPF- ?, Mt Macalister. 
IS""')8^SJ45'''56^E. CaixlvveU Ra.. S0()-900m. 13-16 .Ian. 
1 ^87, SI I (QM S35266). R\RAT\TES: 6 , Mt Macalister. 
Cftrdwcll Ra, 1 000m, sieved litter, 20 Dec. 1986, GBM, 
GT(S35267): 9, 5, Mt Macaiistei; Cardwell Ra. SSQnj, 
pitfall traps 18-20 Dec. 19S6, GBU, GT, SH (S39189); 
26\ 900m, pitfall trap, 18 Dt^ ISSd-M fctit JSSl 
(S39190); (J.pitfeU, 18DeG.198&f*jab.l98?(S39l9t)i 
4dr, 2^ Upper Broadwater Ck,. jS^WS, 145''59'£, 
^rttWeflka., 750m. pitfells, 1 8 Dec. 1986-14 Jan. 1987, 
GBM, GT. SH (S3526S): 27. Mfi Graham, 
18^25'S.1 45^S2'H, Skm N Abcrgowrie. 600-700m, pitfall, 
26 Dec. lL)S6-!7Jan. 1087.Sn(SI4l57); ^^,Kirmina 
Ra,. Main M., W side. i8"I3"S. ]45'M7*E, 700m. pitiall 
traps, lODec. 1986-1 1 .Ictn. 19S7, CiBM, GT, S! I (S30P-^2i; 
V, iCirrama Ra., Mt Smoko luniol]: 18"12'S, 145''46'E. 
600m(S39193); d , Broadwater Park, ^5km NW inuJiam, 
18°12'S, 145°53'E, 500m. pittall, 22 Dec. Jan. 
I9S7, SH (S35271). d\ Walla man Kails, 1S^>6'S, 
I45"4S"F. (ilOw, pirtiills in open forcst, 5-12 Feb. 1996. 
GBM {S35269); J. Minchinbn^okl., 18'^22'S, 146='13'E, 
10m, sieved iiiler. 9 Nov. 1984. VED. GT, J. Gallon 
(S35270H Hinchinbnxik I., GayndaliCk, lOra^pit^ 
8-17 Nov. 1984. \S}}, h Gallpn(S39l94)^ 9, Sam&: 
data (S39 1 9^; C Ifefi- 1^, ^% DC 

DIAGNOSIS. Small spiders (9^ 2.0-3.1). In- 
semination ducts lightly coiled presenting dark 
lateral spots on vetttral surface (c£ all other isppj 



NEW SPIDER GENUS FROM THE WET TROPICS 



TABLE 4. Data matrix. Unknown characters are represented by '?', inapplicable characters by 



Taxa 



10 



Character Number 

20 



30 



40 



Wandella barbarella Gray 033001010 0000000212 000-0-OOUO 000000000- -00 

Dictynidae sp. 120000010 0001001112 1010300100 0012001010 000 

Badumna longinqua (Koch) 010000010 0000100110 0111100100 0013010010 000 

Paramatachia decorata Dalmas 010000010 0000101120 0010100100 0013000010 001 

Desis sp. 110000010 0000202002 1-10300100 0013010010 000 

Quemusia aquilonia Davies 111000010 0000100110 0210100200 0013100010 000 

Jalkaraburra alta Davies 111000010 0000102000 0-10100200 0013100010 000 

Amphinecta milina Forster & Wilton 110000000 001010200? 0-10200100 0011010010 000 

Amaurobiusfenestralis (Stroem) 000000010 0010100110 0010100100 0011010011 000 

Storenosoma terranea Davies 112000001 0010102001 0-10100100 0011010011 000 

Otirasp. 112000001 0010112001 0-10100100 0011010011 000 

Tasmarubrius milvinus (Simon) 112000000 0010102000 0-10101100 0011010111 000 

Procambridgea sp. 100100010 0000100110 0110100100 0011000010 000 

Stiphidionfacetum Simon 011100010 0100100120 0110100100 0012000010 000 

Stiphidion odomatumD^yncs 011100010 0100100120 0110300100 0012000010 000 

Midgee binnaburra Davies 102110000 1000102001 0-10100100 0011000010 000 

Midgee thompsoni Davies 102110000 1000102001 0-10200100 0011000010 000 

Dardurus spinipes Davies 100100010 1000101110 0211100100 0011000010 001 

Manjala plana Davies 110110010 1000100111 0?11100100 0013010010 000 

Malala lubinae DaiVies 100010010 1000102001 0-10100100 0013000010 000 

Kababinaalta Davies 112100110 0100100110 0011100100 0011000040 000 

Carbinea longiscapa Davies 112100110 0100102000 0-11130120 0011000040 000 

Carbinea breviscapa Davies 112100110 0100102000 0-11110120 0011000010 000 

Carbinea wunderlichi Davies 112100110 0100102000 0-11120120 0011000020 000 

Carbinea robertsi DsLvics 112100110 0100102000 0-11120120 0011000030 000 

Malarinamonteithi sp. now. 112100110 0100100110 0011140111 2111000010 000 

Malarinamasseyensis sp, now. 112100110 0100100110 0011140111 0111000010 010 

Malanna collina sp. nov. 112100110 0100100110 0011240110 2111000010 000 

Malarinacardwell sp, my. 112100110 0100100110 0011340110 1111000010 100 



Tibial apophysis short with inturned lateral 
margin, (cf. other spp.) 

DESCRIPTION. Female. CL 1.6, CW 1.2. AL 
1.8, AW 1.3. Legs, I, 6.9; II, 5.5; III, 5.0; IV, 6.8. 
Epigynum (Fig. 4E-H) with tightly coiled 
insemination ducts. Spinnerets (Fig. 5C-E), ALS 
with two major ampullate spigots and about 17 
pirifonn spigots. PMS with a large anterior spigot 
(minor ampullate) and about 12 other spigots 
including 2 (mesa! and posterior) with larger 
shafts (cylindrical) and 3-4 with strobilate shafts 
(paracribellar). PLS with spigots of two sizes. 
Length 2.6-3.4. 

Male. CL 1.4, CW 1 .1 , AL 1 .5, AW 1.1. Legs, I, 
6.4; II, 5.5; III, 4.8; IV, 6.4. Palp (Fig. 41), embolic 
apophysis with all setae about same length; 
plate-like setae reduced in length. Tibial 
apophysis short with inturned edge (Fig. 5F). 
Length 2.9-3.1. 

DISTRIBUTION. Collected from Kirrama/ 
Cardwell Ra., Seaview Ra. and Hinchinbrook I. 
(Fig. 6). 



RELATIONSfflPS OF MALARINA 

A cladistic analysis examined 42 characters 
(Table 3) for relationships of the 4 Malarina spp 
and 25 other taxa. Outgroup comparison was 
with the Australian filistalid spider Wandella 
barbarella Gray and an undescribed Australian 
dictynid. A data matrix for the 29 taxa, names and 
authors given (Table 4), was assembled in 
McClade3.01 (Maddison & Maddison, 1992). 

The data was analysed in PAUP version 3.1.1. 
(Swofford, 1993) and replicated in Hennig 86 
Version 1.5 (Farris, 1988). A heuristic search of 
the data with 10 random -addition sequences and 
TBR branch swapping generated two most 
parsimonious trees differing only in placement of 
M masseyensis — either with M monteithi or 
basal to a clade containmg M. monteithi^ M. 
collina and M cardwell. The preferred tree (Fig. 
7) has length 123, CI = 0.53, CI excluding 
uninformative characters = 0.49, RJ ^ 0.73, RC =■ 
0.39, Characters were mapped in CLADOS 
version 1.2 (Nixon, 1992) with DELTRAN 
optimisation. 



282 



MEMOIRS OF THE QUEENSLAND MUSEUM 



i : ■} b \t> ■ ' . r. r, in 



{-{-Dictynidae .sp. 



rti~i~ 

1 I 

1 



2 14 19 33 

mill 



[) 

1 



-Malala 



3 4 S33 1 ^ 



Midgee binnaburra 



-Midgee thompsoni 



M tD 20 24 



8 \l 33 

> I 
1 



•hAmphinecta 

19 26 37 



3 3» 

2 



a r'^ioretiosoma 
1 I 



HH 

1 



3 27 34 
4ff 



2 

16 n 13 

i-^Jalkaraburra 



1 £> 

n- -Paramatachia 

1 

i 11 33 39 



•+1 

1 



•Aimurobius 



D 1 ( I 



Ll|4. r-BadUma 



++ 



■f^\-MmJala 



2 rhProcambfidgea 

r it 1016 2142 



53 J5 
1 



I 1 

1 



•Stiphidion facetum 

M 

i-Stiphidim adormttwn 

3 
3B 



3 7 21 

+«■ 

2 i 

3 



Kababininae 



r+-t~Mi^^nna massevensis 



I forward chaage 

f forward cliarige with homopiasy 

I reversal with homoplasy 

(j reversal 



25 31 

-H 



16 17 18 25 28 



I \ 1 

Z 



-Malarina monleithi 
-Malarina collina 

li 30 40 

' — ]r\-Molarina cardwell 
-CMrhinea breviscapa 

-Carbinea wunderlichi 

25 38 

^ ^Carbinea robertsi 



FIG. 7. Preferred most parsimonious tree showing tiie cladistic relationships of some Amaurobioidea (branch 
support for nodes in bold type). 



NEW SPIDER GENUS FROM THE WET TROPICS 



283 



Branch support (Bremer, 1994) for the nodes 
on the preferred most parsimonious tree was 
calculated using Autodecay (Eriksson & 
Wikstrom, 1996) and is given in bold type 
beneath each node in Fig. 7. 

CONCLUSIONS 

Wandella and Dictynidae sp. appear as distinct 
from the ingroup which is regarded as the 
superfamily Amaurobioidea. Again this is 
composed of two clades, one including Desis 
(Desidae) and Amphinecta (Amphinectidae), the 
other including Amaurobius (Amaurobiidae), 
Stiphidion (Stiphidiidae) and the metaltellines 
Quemusia and Jalkaraburra. The Kababininae, 
with the addition of Malarina spp. continues to 
form a well supported monophyletic group 
(Davies, 1999). 

The families at the base of the clade are para- 
phyletic therefore the placing of Kababininae 
within any of the existing families remains 
problematic. The group appears to be closest to 
the Stiphidiidae but this is based on a single 
character (feathery hairs) which is also found in 
other genera not represented here. 

While the Kababininae is well supported as is 
the Stiphidiidae, the other families are either 
paraphyletic (eg. Desidae as presently consti- 
tuted) or poorly supported. Support for placing 
the Kababininae within the Stiphidiidae is low. 
Further descriptions and cladistic analyses of the 
Amaurobioidea are necessary to determine 
family relationships and placement of genera. 

ACKNOWLEDGEMENTS 

We are indebted to our colleague Dr G.B. 
Monteith for his and co-workers' collections from 
the Wet Tropics region of north Queensland. 
Since 1993 the field trips have been supported by 
the Wet Tropics Management Authority which 



also supports Kylie Stumkat, SEM technician. 
We thank the Council of the Australian 
Biological Resources Study for funding 
rainforest surveys during which some of this 
material was collected and for the financial 
support of illustrator and co-author, Christine 
Lambkin, who also set up the phylogenetic 
analysis resulting in the cladogram. We are 
grateful for the support of other members of the 
Queensland Museum, particularly Jennifer 
Cannon and Katie Laws for their help in 
preparation of this paper. 

LITERATURE CITED 

BREMER, K. 1994. Branch support and tree stability. 

Cladistics 10: 295-304. 
DAVIES, V.T. 1995. A new spider genus (Araneae: 

Amaurobioidea: Amphinectidae) from the wet 

tropics of Australia. Memoirs of the Queensland 

Museum 38(2): 463-469. 

1999. A new spider genus from north Queensland, 
Australia (Araneae: Amaurobioidea: Kaba- 
bininae). Journal of Arachnology 27: 25-36. 

ERIKSSON, T & WIKSTROM, N. 1996. Autodecay 
version 3.0 (Stockholms universitet Stockholm: 
Sweden). 

FARRIS, J.S. 1988. Hennig 86 Version 1.5. (Port 
Jefferson: New York). 

MADDISON, W.R & MADDISON, D.R. 1992. 
MacClade: analysis of phylogeny and character 
co-evolution, Version 3 documentation. (Sinauer: 
Sunderland). 

NIXON, K.C. 1992. Clados Version 1.2. (L.H. Bailey 
Hortorium, Cornell University: Ithaca). 

PLATNICK, N.I. & SHADAB, M.U. 1975. A revision 
of the spider genus Gnaphosa (Araneae 
Gnaphosidae) in America. Bulletin of the 
American Museum of Natural History 155: 1-16. 

SWOFFORD, D.L. 1993. PAUP. Phylogenetic 
Analysis using Parsimony Version 3.1. (Illinois 
Natural History Survey and Smithsonian 
Institution: Champaign & Washington). 



284 



MEMOIRS OF THE QUEENSLAND MUSEUM 



NEW LOCALITY FOR THE ENDANGERED 
SHEATHTAIL BAT, TAPHOZOUS TROUGHTONI 
TATE, 1952. Memoirs of the Queensland Museum 45(2): 
284. 2000:- Taphozous froughtoni was first described by Tate 
(1952). Subsequently, thetaxon was considered a subspecies 
(McKean & Price, 1967) but was re-established as a flail 
species by Chimimba & Kitchener (1991). It is known from 
only six specimens, four of which are lodged in the AustraUan 
National Wildlife Collections in Canbeira (Chimimba & 
Kitchener, 1991). The three known localities are all in the 
vicinit>' of Mount Isa in NW Queensland. The species has not 
been recorded for 34 years. 

We have identified a specimen of T. troughtoni among 
specimens in the Queensland Museum collection. This 
specimen, registration number QMJ 17608, was originally 
registered as Tadarida australis, but was later re-identified as 
Taphozous australis. We examined QMJ 17608 while 
verilying identifications of Taphozous austraUs specimens 
outside of the species' known distribution (Churchill, 1 998 ). 

QMJ 1 7608 is an adult female. It is in spirit (70% alcohol) 
and in fair condition, although the back of its head has been 
badly damaged. We identified it as an emballonurid by its tail 
protruding from the upper surface of the uropatagium, and in 
the genus Taphozous by its wing pouches and lack of throat 
pouch. We determined QMJ 1 7608 to be T. troughtoni on the 
basis of lengths of its forearm, metacarpal III, and skull 
dimensions. Its forearm was slightly longer than the four 
T troughtoni {16.2 vs 72. 7-75. 6mm) specimens of Chimimba 
& Kitchener (1991), and did not overlap with any other 
Taphozous (Table 1 ). 

It can be difticuh to distinguish between Taphozous 
georgiamfs and T. troughtoni from external characters. 
Examination ofmorphometric data in Chimimba& Kitchener 
( 1 99 1 : 2 1 1 , table 1 b) indicate that the exlemal character with 
the least overlap between these species is the size of the digit 
III metacaipal. The length of this character in the specimen we 
examined was 68.0mm, 0.1mm below the range of T 
troughtoni (68.l-70.2mm, n=4), and 2.1mm above that of 
Taphozous georgi anus (52.7-65.9mm, n=302) (Chimimba & 
Kitchener, 199 f). 

The localit)' recorded for QMJ1760S is 'two miles S.E. 
Cloncurr>- (20^43'S, 140^32'E). It was collected on 15 
September 1 969, although the identit)' of the collector was not 
recorded. The 1971 edition of the Cloncurr>' 1:100,000 map 
(PCC 7056) indicates the probable locality is south of the 
North Western Highway, close to Mount Avarice, with 
quarries and numerous mines within five kilometres. This 
record is around 60-65km to the east of existing locality 
records for T. troughtoni, and so extends its known range. The 
three other locality records for this species are close to each 
other (50kiTi) around Mt Isa. 

An additional specimen and collection locality for this 
species is noteworthy as only six specimens of lliis species 
have been collected from three localities (Tate 1952; 
Chimimba & Kitchener 1991 ). This species has not been seen 
since 1967 (Duncan et al., 1999). The national conservation 
status of T. troughtoni determined by the Action Plan for 
Australian Bats (Duiican et al., 1999) is 'endangered', while 
its conservation status in Queensland is 'endangered' {Nature 
Conservation Act 1994 (and amendments)). A management 
recommendation determined by Duncan et al. ( 1999) is to 
examine museum specimens of Taphozous georgianus from 



TABLE I . Dimensions of morphological characters of 
QMJI7608, T. georgianus and T troughtoni (from 
Chimimba & Kitchener, 199 1). All measurements in mm. 



Character 


QVIJ 17608 


T. georgianus 


T. troughtoni 


Body length 


77.1 


61.6-80.0 


79.4-86.3 


Tail length 


33.8 


22.9-39.4 


31.5-36.9 


Ear length 


24.7 


1 6.5-24.1 


22-4-27.1 


Ear width 


12.1 


1 1.8-25.6 


16.2-19.9 


Tragus length 


7.0 


4-9-8.9 


7.7-9.4 


Forearm length 


76.2 


61.1-73.4 


72.7-75.6 


Tibia length 


32.3 


24.2-30.8 


30.6-31.8 


Pes length 


15.7 


9.8-14.7 


13.8-15.5 


Digit 11 metacarpal 
length 


65.1 






Digit III metacar- 
pal length 


68.0 


52.7-65.9 


68.1-70.2 


Digit III phalanx 1 


23.3 


17.7-23.7 


22.2-24.6 


Digit III phalanx II 


26.1 


19.8-29.1 


25.4-28.4 


Digit IV metacar- 
paflength 


54.S 






Digit IV phalanx I 


15.9 






Digit IV phalanx 11 


11.6 






Digit V metacarpal 
length 


47.3 






Digit V phalanx I 


15.9 






Digit V phalanx 11 


71.8 






Inter upper canine 
distance 


4.5 


3.4-1.5 


4.5^.8 


inter lower canine 
distance 


3.2 


2.6-3.3 


3.2-3.4 


Zygomatic width 


14.7 


12.1-14.4 


14.7-14.9 



northwest Queensland for additional records of T. troughtoni. 
Our findings indicate that examination of presumed 
Taphozous australis specimens may also result in further 
records of 7^ troughtoni. 

Literature Cited 

CHIMIMBA. C.T.& KITCHENER, D.J. 1 991 . A systematic revision 

of Australian Embailonuridae (Mammalia: Chiroptera). 

Records of the Western Australian Museum 15: 203-265. 
CHURCHILL, S. 1998. Australian Bats. (New Holland: Sydney). 
DLINCAN. A., BAKER, G.B. & MONTGOMERY, N. (eds) 1999. 

The Action Plan For Australian Bats. (Environment Australia: 

Canberra). 

McKEAN, .I.L. & PRICE, W.J. 1967. Notes on some Chiroptera from 
Queensland. Australia. Mammalia 31: 101-1 19. 

TATE, G.H.H. 1952. Results of the Archboid Expeditions No. 66. 
Mammals of Cape York Peninsula, with notes on the 
occurrence of rain forest in Queensland. Bulletin of the 
American Museum of Natural History' 98: 563-616. 

A4.P. Rhodes, Queensland Parks and Wildlife Service, PO 
Box J55, Albert Street, Brisbane 4001: A-P. Amey, 
Queensland Museum, PO Box 3300. South Brisbane 4I0I: 
Australia; 10 April 2000. 



GEMMAE OF THE MARCHANTIALES FROM THE WINTON FORMATION 
(MID-CRETACEOUS), EROMANGA BASIN, QUEENSLAND 



MARY E. DETTMANN AND H. TREVOR CLIFFORD 

Dettmann, M.E. & Clifford, H.T 2000 06 30: Gemmae of the Marchantiales from the Winton 
Formation (mid-Cretaceous), Eromanea Basin, Queensland. Memoirs of the Queensland 
Museum 45(2): 285-292. Brisbane. ISSN 0079-8835. 

Hepatophyte gemmae are described from latest Albian sediments of the Winton Formation, 
Eromanga Basin, Queensland. The discoid gemmae are borne on a single-celled stalk and 
midway along each lateral mai'gin there is a shallow notch in which is situated a growing 
point. The gemmae are comparable to those of extant Marchantiales and are referred to 
Marehantites marguerita sp. nov. □ Marchantiales, gemmae, Late Albian, Eromanga Basin, 
Queensland. 

Mary E. Dettmann, Department of Botany. University of Queensland, St Lucia 4072; 
H. Trevor Clifford, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia; 
28 March 2000. 



Hepatophytes are believed to have fonned an 
important component of the mid-Cretaceous 
vegetation of Australia as indicated by the 
widespread, sometimes abundant occurrence of 
diverse hepalic-Iike spores in Albian and Ceno- 
manian sediments (Dettmann, 1994). However, 
apart from the likely affinity of Triporoletes 
Mtchedlishvili to the Marchantiales, ordinal or 
family alliance of the sporae dispersae remains 
speculative. Further support for the presence of 
the Marchantiales in the Albian flora of 
southeastern Australia is provided by the thalli 
taxa, Hepaticites discoides Douglas and H. 
profusus Douglas, both of which are accepted as 
representatives of a possibly extinct, 
marchantialean group (Krassilov & Schuster, 
1984), Spores associated with fertile//, discoides 
conform with the spore genus Triporoletes and its 
junior synonym Rouseisporites Pocock 
(Douglas, 1973). 

In contrast to the common occurrence of 
hepatophyte megafossils, particularly H. 
profusus, at some Albian localities in the Otway 
Basin (Douglas, 1973), there are no records of 
hepatophyte thalli in Albian-Cenomanian mega- 
lloras described from elsewhere in Australia. 
These include the Burrum and Styx compression 
floras (Walkom, 1919); the Winton flora, ktiown 
from impressions and permineralised cones and 
foliage taken froin outcrops (McLoughlin et al., 
1995; and references cited therein); and recently 
described compressions and cuticles recovered 
from core material of subsurface strata (Pole, 
1999; 2000, Pole & Douglas, 1999). During 
palynological processing of a core from the 
Winton Formation in GSQ Thargomindah No. 3. 



numerous mesofossils including megaspores, 
fern sporangia, and discoid hepatophyte gemmae 
were recovered. This account details the gemmae 
and illustrates associated fern sporangia. 

MATERIAL AND METHODS 

The gemmae were isolated froin a siltstone 
intersected at 163.5m in GSQ Thargomindah No. 
3, a continuously cored stratigraphic borehole 
drilled by the Geological Survey of Queensland 
at 27^16^S, 142°55'E, 120km NW of 
Thargomindah within the Eromanga Basin, SW 
Queensland (Figs 1, 2). A routine check of 
organic matter extracted from the sediment after 
treatment with 50% hydrofluoric acid followed 
by thorough washing in distilled water revealed 
the presence of numerous small discoid plant 
fossils up to 440|xm in diameter. These were 
picked from the residue and transfeired to small 
petri dishes prior to mounting in glycerine jelly 
on glass microscope slides. No further chemical 
treatment was required, and indeed mild ox- 
idation in dilute nitric acid resulted in destruction 
of the disc-shaped fossils. After recognising that 
the discs represented hepatophyte gemmae, a 
thorough search was undertaken for any 
associated hepatophyte tissues; none was foimd, 
except for occasional hepatophyte-Iike dispersed 
spores {Triporoletes reticiilatus (Pocock) 
Playford, T simplex (Cookson & Dettmann) 
Playford, and T. radiatus (Dettmaim) Playford). 
Other plant meso/microfossils represented in the 
residue included fern sporangia, woody tissues 
and palynomoiphs. Fern sporangia were picked 
from unoxidised portions of the residue and 
mounted either in glycerine jelly for light 



286 



MEMOIRS OF THE QUEENSLAND MUSEUM 




EROMANGA 
BASIN 



GSQ Thargomind^h - 3 



• i;hargomindah Brisbane - ^ 



CAINOZOIC 













deplh (m) 



100 



Winton 
Formation 




163.5m 



Mackunda 
Formation 



Allaru 
Mudstone 



200 



300 



400 



sandstone and 
silty sands 



mudstones/ 
shale 



FIG. 2. Stratigraphic sequence in GSQ Thargomindah 
No. 3, and sampling horizon. 



— - - basin margin 

FIG. 1. Map of Australia showing location of GSQ 
Thargomindah No. 3 and Eromanga Basin. 

microscope examination or on stubs and sputter 
coated with gold for scanning electron micro- 
scope analysis. Palynomorphs were extracted 
after treatment with nitric acid for 2 minutes, 
followed by thorough washing in distilled water, 
brief immersion in 1% ammonium hydroxide and 
further washing in distilled water prior to 
mounting for light microscope analyses. 

The palynoflora contained in the sediment 
indicates assignment to the upper part of the 
Phimopollenites pannosus spore-pollen Zone (of 
Helby et al., 1987) and thus a latest Albian age. 

Gemmae of living Marchantia berteroana 
Lehm. et Lindenb., Liimilaria cruciata (L.) Dum. 
and Neohodgsonia mirabilis (Perss.)Perss. were 
examined after clearing in a mixture of glacial 
acetic acid and hydrogen peroxide in proportions 
7: 1 to remove chlorophyll and cell contents. The 
gemmae were then washed in distilled water and 
mounted in glycerinejelly onmicroscope slides. 

Type and other figured specimens are lodged in 
the Queensland Museum, Brisbane. Registered 
numbers of that institution are designated in 
Table 1. 



SYSTEMATIC PALAEONTOLOGY 

HEPATOPHYTA 
MARCHANTIALES 

Marchantites Brongn., 1849 
emend. Walton, 1925 

TYPE SPECIES . Marchantites sezanmnsis Brongn., 1 849. 

Marchantites marguerita sp. nov. 
(Fig. 3A-K) 

ETYMOLOGY For the late Margaret Derham, beloved 
sister and friend of MED. 

MATERIAL. HOLOTYPE: QMF50093. Fig. 3A-C. 
Gemma discoid, 430|xm long, 350|xm greatest width, with 
a stalk scar at one end, and a notch on opposite sides on the 
periphery situated lateral to the stalk. Cells polygonal, 
30-40|jL in diameter, with anticlinal walls up to 6|xm high. 

DIAGNOSIS. Gemmae discoid, 1-2 cells thick 
and sometimes with a short, one-celled stalk up to 
60|jLm long and 60|xm wide. In outline each 
gemmae is bilaterally symmetrical about the 
vertical axis. A pair of shallow notches occurs 
opposite to each other on the perimeter of each 
gemmae midway between the apex and stalk. 
Cells adjacent to stalk elongate (up to 80|jLm long, 
30-40|jLm wide), elsewhere isodiametric, 
pentagonal to hexagonal, 25-40|jLm in diameter 



MARCHANTIALHS FROM THE tROMANOA BASJN 



287 




FIG. 3. Micrographs of A/i/rr/w/?///t\v marguerita sp. nov.: A-C\ holoL>pe. A. X 100;B. basa! cells at site ofsialk 
atlachment x300: C, cells in region oi" notch, x 300; whole specimen, X200, and detail of notch cells, 
X 300; F,G specimen with stalk attached, x 100, and detail of stalk ceil, X 150; H, i. specimen X 150 and 
X 100: J. K. specimen, x 100. and detail of cells, x 300. 



andwilh anticlinal walls2-6^jim high, but grading DIMENSIONS, (longituditjal X latesral dimen- 
to 20|jLni in dianiet«^ a« the g^oiVitig p6ttit& ^Oiis) 200-(320)-440|j,io X 160-(270)-400|Jim 
centred TIT each latwal notch, (20 specimens). 



288 



MEMOIRS OF THE QUEENSLAND MUSEUM 



A B 



- ■■■\ - y 



n 



H 




D 




FIG 4. A-E, Gemmae oiMarchantia herteroana Lehm. et Lindenb; A, X 1 00; B, C, detail of notch and peripheral 
cells, X 300; D,E, specimen with stalk cell, X 100 and detail of stalk, x XSO.YA.GQmm'dQoiLumdwiacruciata 
(L.) Dmn; F-H, specimen, x 100, and detail of cells in central and basal regions, x300; I, specimen, x 100. 

TYPE LOCALITY. GSQ Thargomindah 3, morphologically consonant with gemmae of 

163.5m; \xp}?Qr P. pannosus Zone, latest Albian. Marchantia L. (Marchantiaceae) and Lumdaria 

REMARKS AND COMPARISON. In Adanson (Lunulariaceae). Cells of Marchantites 

possessing a single-celled stalk and peripheral ^'-^-f/'^wenYa are more similar in size {25-40fim in 

lateral notches in each ofwhich is centred a small diameter) to those of gemmae ot Marchantia 

growing point, the discoid fossils are berteroana (cells 25-40fxm in diameter; Fig. 



MARCHANTIALES FROM THE EROMANGA BASIN 289 



4A-E) than those oi Marchautia polymorpha L. 
(Smith, 1955: fig. 30; 15-30|xm in diameter) and 
Lumilaria cruciata (cells 15-30|xm; Fig. 4F-I). 
The monospecific Neohodgsonia (N. mirabilis) 
H. Persson, (Marchantiaceae) also has 
disc-shaped gemmae, but they differ from those 
examined of Marchantia and Limularia in 
possessing only one growing point associated 
with a lateral or subapical notch. Plate-like 
gemmae occur in several extant taxa of the 
Metzgeriales, but in these there is only one 
growing point, which is situated at the margin 
opposite to the stalk (Watson 1964). 

The likely Marchantiales origin of the fossil 
gemmae may argue for assignment to 
Marchantiolites Lunblad, also demonstrated to 
be consistent with the Marchantiales. However, 
Marchafitiolites is based on thalli with rhizoids 
on the undersurface and air pores on the upper 
surface, and is thus inappropriate for the gemmae 
described here. Pending recovery of the gemmae 
in organic association with hepatophyte thalli, 
the fossils are included in the broader category 
Marchantites. 

DISCUSSION. Because of their firm thalli and 
preference for growing on mineral soil adjacent 
to stream banks, members of the Marchantiales 
are likely candidates for burial and subsequent 
fossilisation. Indeed from the Triassic onwards 
marchantioid-type thalli are well known 
(Krassilov & Schuster, 1984). It is therefore 
surprising that Marchantia-WkQ gemmae have 
not been previously recognised. 

The similarity of Marchantites marguerita to 
gemmae of extant Marchantia and Limularia 
suggests that growth habitats of the Winton 
hepatophytes were within the range of those 
occupied by the extant genera, both of which are 
restricted to temperate climates. The 
Thargomindah region of the Eromanga Basin 
was situated at ~55°S diu-ing the latest Albian. 
Palaeotemperature data are lacking but those 
deduced from belemnites and bivalves from the 
underlying marine sequence indicate sea water 
temperatui-es of 1 2- 1 6"C (Dettmann et al., 1 992 ). 
Temperatures adduced from Global Grossplots 
are near 15°C (Frakes, 1997) and from other 
sources approximate 1 0°C (Frakes, 1 997, Fig. 4). 
Today, the 15°C MAT isotherm passes through 



southern New South Wales/northern Victoria at 
latitudes close to 35°S and the 10°C MAT 
isothenn to the south of Tasmania (Anon, 1988). 

Associated with the fossil gemmae are 
numerous fern sporangia, one type possibly 
osmundaceous (Fig. 5A-D; apical annulus and 
containing in situ Osmundacidites cf wellmanii 
Couper), and another w ith a vertical annulus and 
containing Cyalhidites minor Couper (Fig. 5E-I). 
Also represented are lycophytic and hydropter- 
idean megaspores and a restricted spore-pollen 
flora dominated by tllicean spores referable to 
Ruff or diaspora Dettmann 8l Clifford 
(Schizaeaceae). Baculatisporites Tliomson & Ptlug 
and Osnnmdacidites Couper (Osmundaceae), and 
Cyalhidites Couper. Hepatic spores {Triporoletes 
reticuIaTus, T. simplex and T. radiatus)^ 
gymnosperm {Araucariacites Cookson ex 
Couper, Podocarpidites Cookson ex Couper, 
Microcachyidites Cookson ex Couper) and 
dicotyledonous angiosperm pollen (Phimo- 
pollenites Dettmann) occur in low frequencies. 

ACKNOWEEDGEMENTS 

Grateful acknowledgment is extended to the 
Queensland Department of Mines and Energy for 
enabling sampling of core material; Dr Mike 
Pole, Department of Botany, University of 
Queensland for assistance with sample 
collection; Dr David Bedford, Royal Tasmanian 
Botanical Gardens for provision of material of 
Marchantia herteroana\ and Dr Patrick 
Brownsey, Museum of New Zealand - Te Papa, 
Tongarewa, New Zealand for provision of 
material of Neohodgsonia mirabilis. We 
acknowledge helpfi^d reviewers comments of Dr 
D. Cantrill and Dr J. Douglas. MED extends 
sincere thanks to Dr P Jell Acting Director, 
Queensland Museum for providing facilities. Tlie 
research was supported by Australian Research 
Council Grant A 1953 1 899 to MED and Professor 
George Stewart. 

LITERATURE CITED 

ANON. 1988. Climatic averages of Australia, (Bureau 

of Meterologv: Canberra). 
DETTMANN, M.E. 1994. Cretaceous vegetation: the 

microfossil record. Pp. 143-170. In Hill, R.S. (ed.) 

History of the Australian vegetation. (Cambridge 

University Press: Cambridge). 



FIG. 5. A, B, Osmundaceous sporangia, X 150. C, D, Osmundacidites wellmanii Couper, X 750; C, spore 
associated with sporangium, D, dispersed spore. E, Baculatisporites comaumensis (Cookson) Potonie, 
dispersed spore X 750. F-I, Fern sporangia with vertical annulus; F, x 400, G, with in situ spores, X 200, H, I, 
x250. J, Cyathidites minor Couper, spore from sporangium, x750. K, Triporoletes simplex (Cookson & 
Dettmann) Playford, dispersed spore, x 750. 



MARCHANTIALES FROM THE EROMANGA BASIN 291 



TABLE 1. Register of figured specimens. * denotes holotype. 



Taxon/Fig. No. 


Slide 


Co-ordinates (England Finder) 


Registered No. 


Gemmae 


Liinularia cruciata 


Fig. 4F-H 


LUN/2 


K28/1 


QMF50110 


Fig. 41 


LUN/l 


L34/2 


QMF50111 


Marchantia berteroana 


Fig. 4A-C 


MARyi 


Q41 


QMF50112 


Fig. 4D,E 


MAR/1 


K26/3-4 


QMF50113 


Marchantites marguerita 


Fig. 3A-C * 


THA 163.5/A2 


Mil 


QMF50093 


Fig. 3D,E 


THA 163.5/A5 


E37/1 


QMF50094 


Fig. 3F,G 


THA 163.5 /All 


052/4 


QMF50095 


Fig. 3H 


THA 163.5/A8 


G29 


QMF50096 


Fig. 31 


THA 163.5/A2 


CI 6/4 


QMF50097 


Fig. 3J,K 


THA 163.5/Al 


K41/1 


QMF50098 


Sporangia 


Osmundaceous sporangia 


Fig. 5A 


THA 163.5/A7 


P38 


QMF50099 


Fig. 5B 


THA 163.5/K 


F29/4 


QMF50100 


Sporangia with vertical annulus 


Fig. 5F 


THA 163.3/A6 


J33/3 


QMF50101 


Fig. 5G 


THA 163.5/A3 


H27 


QMF50102 


Fig. 5H 


THA 163.5/A16 


L38/1 


QMF50103 


Fig. 51 


THA 163.5/A16 


M37 


QMF50104 


Spores 


Baculatisporites comaumensis 


Fig. 5E 


THA 163.5/2 


F31 


QMF50105 


Cyathidites minor 


Fig. 5J 


THA 163.5/2 


K50/2 


QMF50106 


Osmundacidites weUmanii 


Fig. 5C 


THA 163.5/2 


J45/3 


QMF50107 


Fig. 5D 


THA 163.5/2 


Q46/4 


QMF5O108 


Triporoletes simplex 


Fig. 5K 


THA 163.5/2 


J43 


QMF50109 



zoic. Association of Australasian Palaeontologists 
Memoir 4: 1-94. 

KRASSILOV, V. & SCHUSTER, R.M. 1984. 
Palaeozoic and Mesozoic fossils. Pp. 1172-1193. 
In Schuster, R.M. (ed.) New manual of bryology 
Vol. 2. (Hattori Botanical Laboratory, Nichinan: 
Myazuki, Japan). 

McLOUGHLIN, S., DRINNAN, A.N. & ROZEFELDS, 
A.C. 1995. A Cenomanian flora from the Winton 
Formation, Eromanga Basin, Queensland, 
Australia. Memoirs of the Queensland Museum 
38:273-313. 

POLE, M. 1999. Latest Albian-Cenomanian mono- 
cotyledonous leaves from Australia. Botanical 
Journal of the Linnean Society 129: 177-186. 



DETTMANN, M.E., MOLNAR, R.E., DOUGLAS, 
J.G, BURGER, D., FIELDING, C., CLIFFORD, 
H.T, FRANCIS, J., JELL, P, RICH, T, WADE, 
M., RICH, RV, PLEDGE, N., KEMP, A. & 
ROZEFELDS, A. 1992. Australian Cretaceous 
terrestrial faunas and floras: biostratigraphlc and 
biogeographic implications. Cretaceous Research 
13:207-262. 

DOUGLAS, J.G 1973. The Mesozoic floras of Victoria. 
3. Geological Survey of Victoria Memoir 29: 
1-185. 

FRAKES, L.A. 1 997. Grossplots: a method for estimat- 
ing the temperature state of the Earth and 
Australia, Cretaceous to Middle Miocene. 
Australian Journal of Botany 45: 369-372. 

HELBY, R., MORGAN, R. & PARTRIDGE, A. 1987. 
A palynological zonation of the Australian Meso- 



292 



MEMOIRS OF THE QUEENSLAND MUSEUM 



2000. Mid Cretaceous conifers from the Eromanga 
Basin, Australia. Australian Systematic Botany 
13: 153-197. 

POLE, M.S. & DOUGLAS, J.G. 1999. Bennettitales, 
Cycadales and Ginkgoales from the mid 
Cretaceous of the Eromanga Basin, Queensland, 
Australia. Cretaceous Research 20: 523-538. 



SMITH, G.M. 1955. Cryptogamic botany, Vol.2, 
Bryophytes and Pteridophytes. (McGraw Hill: 
New York). 

WALKOM, A.B, 1919. Mesozoic floras of Queensland, 
Parts 3 and 4. The Hotm of the Bumim and Styx 
River Series. Queensland Geological Survey 
Publication 263: 1-76. 

WATSON, E.V. 1964. The structure and life of Bryo- 
phytes. (Hutchinson; London). 



CLIFEOKI> 8. FRnmND DAWM W. mm 

KrilliuC.Bt & Frith. D. W. 2000 06 30: Bower sysicin and slruclures ol ihe Golden liowcrbird. 
Pno^otfy^'a i\}SwtQHktmi (Wlonorhyqchitl^eK Mmoirs qfihc Queemktttd Museum 45(2): 

Wo expmmcd SQGoIdcit BowerbW; f^ipnc«iii)fQ*icimiom(tm, bo^yersi1fc8, Ihvtilvihg a total 
of 98 mailt (jdepcnAlejl) hnvcr^ctulcs tiuring W8-t997.0nly one main bower siructtire 
Wn5 d^Vti^^acttVcl^ ',!ltany bOVwrsltii during any one season. Bower sites were iradiliotuil 
(n = 49) or nidimeniary (n - 11). Traditional sites were dispel led spatially throughtml 
sellable Jopopaphyat an average of one pei 4.2ha, and at a iiiLan nearest neighboui* distfliKC 
of 1 5l ± 44m (rt= VZ). Eighly'tbur per ccni o\"25 tradiiional siics were regularly aitendcd 
cnnKeculWcly for 20 seasons. RudiniciiUuA sites \scrc locaicd ^8 t 36m Trom iniilitional 
shcs and were raj'ely aelivc Ibr more than i\\o seasons. Tradiiional bouoi:- con r 
single (36"o) or iwo (64Vn) lowers when tii ^t found; 1-1 of ilic single U)\\ei iHiov L i -. laier 
became two tower structures. Eaeh bower had a bower pereh of woodv vine., near- lion. ^onial 
li\ ijig sapling. fallL*n dead hnineh or tree root averaging 4.6 - 6.5eni diaitjeler. \\ here ilicy 
abutted the bower pereh. unver sticlvS were aligned lightly into a plallorni(;.). upon whieii 
decorations, tApicaiU gre\ ish-gi'een iiehcn I'snca sp, and creamy-wiiitc suod puds of 
Me/icopc hroiidbeniinnu, were placed. Mean minimum age of a liadillonal bower \\ a,> 'J. 6 ± 
(}3 yearb (n = 48), at a mean of two per site over time. Six such bowers were attended tor 20 
seasons. Mean distance of a new traditional bower structure from the replaced t)ne was 14.1 
± I2.7ii5. Nc^v main bower structures started as small single Drlxireal conical or 
niaypole-shaped slruclures. k tooli two lo tlires: seasons fgrtheni to rcaeh full si^c. l ourleen 
arboreal lowers of main bowers subsequently betiaoiBt^tcstrial, because sticks aeeumii|Bt?d 
beneath theiin. Small arboreal and terrestrial ititisl^G^^cnvei' structuj'cs. built at aitipaflr 
distance of 5.4 - 4.2m (torn main bower stmctutfeS^CKpi^linfi^fo^ Itje bases iue^ 
main one; suggesting a function of subsidiary strucVure^/WecdiiehKltsdlArTvii^^ 
iWi^ shfiyslc arc Jipt ©pnsemtive in this bowcrbird, the platform area(s) upon which 
^CCOVallOftB ^ft^^lactfd, art conservative in being ftpccifieally Iwatcd, better constructed 
and in being decorated. The significance of boWer fbnn ana adull m:ile plumage in the 
Golden Bowerbird ate discussed., d Golden H^^erbUxi, Phanot/t/ra ne^'iOfUmcti 

C/iJftin/ a j-'rHh unci Dmvn \V Friifh, Bonontry Racurch Fi>llo\\ s of (he Oitv^liitljd 

Mn-icum 'PnontyJuri] ', PO Box 5SL Mdlamh V'S'.S'.r Ausirulia, J7 ScpicTnfK'rl<W 



The Golden Bow erbittl, Priurmimrti ffnyy/otpamt, 
reprcscntB a distthctive monotypk genus ctidcmic 

10 the Atherton Region of the Australian Wcl 
Tropics, tropical northeast Oucensland, above 
680 m asl (Blakcrs et aJ., 1984; Ni\ Swii^xi, 
lyyi). Tliis species, the smallest bowerbird (25cin 
in'-letigth and averaging 75g), is strikingly 
sexually dimorphic. Adull-pliimaged males are 
predominantly brilliant yellow on their tindcr- 
parts. brownish olive above with a small bright 
yellow crest on the central crown, and a larger 
yldlovs nape patch. Females and immature males 
are pale grey below and brownish olive above. 

The Golden Bowerbird is one of 1 6, of the total 
19, "bowerbird species (Ptilonorhyndvidae) that 
hasapdygNaicnis mating system, in w hich males 
urt? pcomiscuoi^ and fetaude^^ build and attend 
nests alone* M3le$ <b«Sld Jtla|ge'mi^)K>w^ (Figi 



1 ) of the 'maypole' lyp^. as do the four 
hiwerbfrdfttaflhelSfWOC^ 

This h quite unlike the cleared 'court' t)i'SrenO' 
poetics, the "mat' of accumulated lern fronds' of 
ArchhttUIiu. or the stick or grass 'a\cniic' type 
bowers of Fnkmorliynchus, Chlamydcra ;(nd 
Sericiilus spp. (Marshall, 1954; Ciilliard, 19(59; 
Cooper I orshaw, 1977; Borgia, 1986; Frith, 
I9H9; Frith et al., 1^94, i996a,h; Ftilh 8l Frith, 
1W3. i^W. lW;Donaghcy, 19SK WHi). 
Bowers ol most bowerbird species require re- 
building or major reftirKshincnt within and 
between each display season, but maypole 
bowers of Golden Bowerbirds. and of some of tlie 
closely related gardcnci bowerbirds, petsisi vear 
toycar(Priiett-Jones&Fruett-.Iones, I'>82. 1983). 

The Golden was the last bowerbird to be 



294 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FlCl. 1 . A tradilional twin tower bower and its adult male tiolden Bowerbird owner. Note where lower slicks meet 
the bower perch (black in Ihis picture) Ihey are more ski 1 1 fully placed and aligned into a discrete 'platform', than 
are slicks ofthe rest of the slruclure. Decorations are placed on and adjacent to these 'plaifomis' and those seen 
here are beard lichen Usnea sp., seed pods of Melicope broadbentiaiui ajid the creamy white llowers of 
Darlingia darlingiana (one in bird's bill). 



Chisholm & ChafTer (1956). Bowers were first 
described by Broadbent (in Campbell, 1900), 
Day (in North, 1904); Broadbent, 1902; North, 
1909; Sharp (in North, 1914) and De Vis (in 
Mathews, 1926). Photographs of a bower 
appeared in Jackson (1909), but it was not until 
much later that more bowers were described 
(Bourke & Austin, 1947; Warham, 1^62; 
Chishoim & ChalTer. 1956; Chaffcr, 1958. 
Chisholm, 1957, 1963; Gilliard, 1969; Marshall. 
1954). Bowers typically consist of two stick 
towers, which may or may not be ofeqtial height, 
ora bulky and irregularly-shaped single massif of 
sticks with a bowser perch protruding Irom one 
side. Structures vary considerably (Frith, 1989). 
Each lower is built upon and around a supporting 
central sapling(s) or tree. Twin tow er bow ers are 
up to 1 m apart and are interconnected by a living 
or dead, arboreal or terrestrial, horizontal or 



near-horizontal, perch. Bower decorations are 
placed on the more neatly-aligned tower sticks 
adjacent to and on the bow er perch. These include: 
greyish-green lichen Usnea sp., creamy-while 
seed pods Melicope hroadhenticma. and w hilish 
flowers of several plant species (Chisholm 8l 
ChalTer, 1956; Chaffer, 1958; 1984; Warham, 
1 962, Frith & Frith, 2000a). Several other, small- 
er, stick subsidiary bovver structures Cgunyahs' 
ol'Broadbenl, in Campbell, 1 900), are built close 
to the main (decorated) bower strucuire. Bower 
building/maintenance/decoration reaches a peak 
during the display season, from late August- 
December on the Paluma Range. Such activity 
declines during the heavier wet season rains of 
Jajiuai7 and/or February, and when birds are 
moulting. Renewed, post-moull, activity com- 
mences in mid-March and April (Frith &. Frith, 
2000a,b). 



BOWERS OF THE GOLDEN BOWERBTRD 



295 




FIG. 2. Dispersion of 41 traditional bower sites of male Golden Bowerbirds: 12iti study area 1, 10 in study area 2, 
and 19 in adjacent extralimita! areas. Each 50ha study area measured 1 X 0.5km. Those numbered are the 25 
traditional sites examined seasonally (S78-S97; see Table 4). Note: single lines show creek systems, double 
parallel lines represent the dirt road from Paluma Township (entering at bottom) to Paluma Dam (to north) with 
a side track through SA 1 ; the dotted line shows a snig-track through SA2 forest. 



Bowerbird studies are numerous, but few deal 
systematically with variation in bower/court 
structures/sizes (as opposed to bower decorat- 
ions) or provide comparative measurements of 
them. Exceptions are those of Borgia (1985 ) for 
the Satin, Diamond (1987) for Vogelkop, 
Amblyornis inornatus, Lenz (1993) for Regent, 
Sericiihis chry^socephalus. Frith & Frith (1994) 
for the Tooth-billed, Scenopoeetes dentirostris. 
Frith et al. (1996a) for Archbold's, Archboldia 
papuensis, and Frith et al. (1996b) for Great 
Bowerbirds, Chlamydera nuchalis. The present 
study, carried out during 1978- 1 997, provides the 
first detailed information on variation in the 
structure and size of bowers of Golden Bower- 
birds. It includes information on bower site 
location and dispersion, bower site constancy, 
bower age and bower building. Data on male 
seasonal activities at bowers, including attend- 
ance levels, bower maintenance, vocalisations, 
displays, decoration theft, and home ranges are 
presented elsewhere (Frith & Frith, 2000a,b), as 



will be data on bower ownership, male survival 
and home ranges (Frith & Frith, unpubl. data). 

METHODS 

STUDY AREA. This study was perfonned in 
upland tropical rainforest, classified as simple 
notophyll vine forest (Tracey, 1982), at about 
850m asl, 7km from Paluma Township on the 
Paluma Range, NE Queensland. The main 50ha 
study area (SAl, at 19°00'S,146°10'E)measured 
1 X 0.5km and was permanently gridded with 
metal stakes (see also Frith & Frith 1 994, 1995). 
A narrow dirt road bisected the length of the 
broad main ridge line of SAl (Figs 2, 3). To the 
north of this road was a broad flattish ridge 
30-50m wide and 600m long; with a discrete hill 
from which a slope, dissected by gullies, fell 
steeply down to a perennial creek. To the south 
the ridge was flatter and wider (240m); 
interspersed with patches of Calamus-domm^XQd 
undergrowth and a system of creeks, except at the 
western end where it rose to a ridge and another 



296 



MEMOIRS OF THE QUEENSLAND MUSEUM 




+ 045 



+ •♦■ ,(10m) #21 (15m) ; : #29 (38m) 

FIG. 3. Dispersion of 15 traditional (•) and 10 rudimentar>' (+) bower sites of male Golden Bowerbirds in study 
area 1 and outside (distance indicated in figure) its perimeter. Note: one rectangular quadrat = 2ha; number in 
top right hand comer of each = topographic type predominant (>75%) in that quadrat. Faint lines indicate 
location of seasonal gullies. 



hill. Beyond this area the terrain dropped away 
steeply. A second hill-side study area (S A2) of the 
same size was not gridded. It was contiguous with 
SAl and extended northwestwards up a hill to 
950m asl. An old forestry snig-track bisecting a 
narrow ridge provided access up this hill (Fig. 2). 
Both study areas were searched systematically 
for bowers by CBF during August 1 978-February 
1981 for a total of 975h, besides innumerable 
unrecorded hours of random searching diu-ingthe 
course of this and other bowerbird studies (Frith 
& Frith, 1994, 1995, 1998). Extralimital areas 
along tracks and ridges for a distance of up to 2km 
beyond SAl and SA2 were also casually 
searched, as were other areas around Birthday 
Creek Falls and Paluma Dam. 

DEFINITIONS. ^Site' describes the location of 
any active bower found, and any replacement 
bower(s) built subsequently during the course of 
the study. A traditional bower site was one 
attended for at least two seasons (Frith & Frith, 
1994). A traditional bower was a large and 
well-established (single or twin tower) structure 
that was regularly attended, maintained, and 
decorated, throughout subsequent seasons by its 
traditional adult male owner. In a few instances, 
after the disappearance of a long-term traditional 
adult male owner, a traditional bower site was 
irregularly attended by immature (female- 



plumaged) males, who either maintained the 
existing traditional bower or built a new 
rudimentary structure at that site (Frith & Frith, 
2000b). Such a rudimentary bower subsequently 
became larger, and a traditional one, once an 
adult male again attended the site regularly. 

A rudimentary bower site was one established 
near a traditional site by the construction of a 
rudimentary bower. Such bowers were poorly 
constructed, maintained, decorated and irreg- 
ularly attended for only a few days/weeks each 
season, by immature males. We use regularly 
attended to imply full-time seasonal attendance 
by traditional owners at traditional bowers, and 
irregularly attended' to imply part-time seasonal 
attendance by immature males at traditional or 
rudimentary bowers. We refer to a display season 
by the year in which it started (S78, S79 etc). 

BOWER SITES, THEIR LOCATION AND 
DISPERSION. We located 60 bower sites and 98 
main bower structures (Table 1 ). Each bower site 
was plotted and each bower numbered (site 
number followed by suffix a, b, etc., for every 
bower built at a site) and tagged with a scored 
aluminium label on a tree supporting a bower 
tower. 

The possible influence of diftering topography 
upon bower structure and dispersion of 12 
traditional bower sites in SAl was examined. 



BOWERS OF THE GOLDEN BOWERBIRD 



297 



TABLE 1 . Number of bower sites, bower types (traditional or rudimentary) and bower structure of male Golden 
Bowerbirds and the type of bower structure (single or twin tower) when first found on the Paluma Range, north 
Queensland. * = at least 14 became twin tower bowers during the study. 





Total number 


Number of old bowers (before 1978) 


Number of new bowers (1 978-1997) 


Sites 


Bowers 


Single tower 


Twin towers 


Total 


Single tower * 


Twin towers 


Total 


Traditional 


49 


86 


I 


11 


12 


30 


44 


74 


Rudimentary 


11 


!2 


3 





3 


8 


1 


9 




60 


98 


4 


11 


15 


38 


45 


83 



Various quadrat grid sizes were uniformly 
applied to a topographic map of the study area to 
find the most suitable grid size that resulted in 
each quadrat containing >75% of a given 
topographic type (see Frith & Frith, 1995). This 
proved to be a grid of 25 quadrats each of 2ha, or 
200 X 100m. Each quadrat was assigned to the 
topographic type predominating (Fig. 3) as 
follows: 1 ) very steep slopes of >40° dissected by 
gullies (8ha); 2) steep hill slopes of 20-40° (4ha); 
3 ) hill tops with 1 0-20'' slopes (4ha); 4) gentle hill 
slopes of 5-10'' (6ha); 5) ridge-side with <10° 
slopes (8ha); 6) open flat or <5° sloping areas 
(6ha); 7) disturbed flat areas with dense under- 
storey dominated by Calamus (4ha); 8) flattish to 
<5° sloping areas dominated by creek systems 
and dense understorey (lOha). To test whether 
dispersion of bower sites was random, the 
numbers of sites per quadrat was compared to 
expected Poisson distributions. Coefficients of 
dispersion (CD: variance to mean ratio) were 
calculated as a quantitative description of 
dispersion. This method is based on the variance 
being equal to the mean in the Poisson 
distribufion. Variance to mean values of 1.0 
imply random, >1 implies clumped, and <1 
implies a regular or spatially unifonn distribution 
(Sokal and Rohlf, 1969). The significance level 
of an obsei*ved deviation of the CD. from 1 .0 was 
determined by a t-test (n-1, one-tailed), the 
t-value being calculated by dividing the differ- 
ence between the CD. and 1.0 by the standard 
error of the deviation. 

Mean distances between traditional bowers at 
12 traditional sites in SAI and at three other sites 
just outside its perimeter (Fig. 3) were estimated 
in two ways. First, nearest neighbour distances 
(NND) between sites were analysed using the 
method of Clark & Evans (1 954). In this method: 
when two sites are closer together than they are to 
any other ones then the same distance is included 
twice. Secondly, although bower sites were not 
arranged linearly in this area, we estimated the 
mean inter-bower distance. This involved taking 



the measurement from one bower to the next 
closest and so on throughout the whole 50ha. This 
allowed us to compare linear inter-bower 
distances with those presented in other bowerbird 
sUidies. 

BOWER CHARACTERISTICS. Seventy-seven 
traditional bowers were measured and photo- 
graphed. The following measurements were 
taken: height and base circumference of each 
tower and, in twin tower bowers, the distances 
between tower bases and apices; the type, axis 
direcfion, height and diameter of the bower perch 
and, in twin tower bowers, the length of perch 
exposed between the platforms of the towers; the 
number and girth at breast height (gabh) of 
saplings and trees incorporated into each tower. 
Number and size of associated arboreal and 
terrestrial subsidiary structures were measured 
and their distances relative to the main bower 
perch plotted. Means are given ± one standard 
deviafion. 

To give an indication of relative bower size we 
estimated bower volume by multiplying tower 
height with base circumference. Spearman rank 
correlafion (one-tailed test conected forties) was 
applied to test whether there was a correlation 
between bower size (= volume) and situation (= 
degree of slope) at 42 traditional sites. When we 
measured two bowers at a site (n = 6) we took the 
mean value of each measurement. 

BOWER SITE CONSTANCY, BOWER AGE, 
BOWER BUILDING AND STRUCTURAL 
CHANGES. To provide data on site constancy 
we examined 25 traduional bower sites from 
S78-S97 but excluding S9 1 , S94 and S96 (Fig. 1 ). 
Our absences during these latter seasons did not 
affect our results as all sites save one (site 27) 
remained actively attended by birds during the 
subsequent season(s). We omitted season 91 be- 
cause it was excessively dry, bowers were seldom 
attended and were poorly decorated. Rainfall 
typically averaged 259mm (S78-S90) for 
September-November; but in S91 only 94mm of 
rain fell, mostly after 12 November. 



298 



MLMUIRS Ob THE QUEENSLAND MUSEUM 




FIG. 4, Shapes and si/es of single tovs cr hos\ ci s. ( scales ~- a one metre stick marked every 1 0cm or CBF ( 1 80cin 
tall) or DWF ( 162cm tall) in piciure). /\, houcr 37a: a compact single tower vvilli a cur\'ed vine bower perch 
(right) with small terrestrial suhsidian deft in background). April 1979. B, bower 23a: a bulky single 
irrvgular-shaped massirwiih a sloping living-sapling bower perch (right). Apnl 79. "Phe llnv irregularly-placed 
sticks Lo ihe righl end ot the bow er perch never hecaine a second tower and the bovver changed little in shape or 
si/e over six .seasons. C, bower 22a: an amorphous iliree-peaked massif with a rotten bovver perch on the ground 
( left ofphotograph ). Apri 1 1 979 /Fhc owner had replaced this bower with a new one by September 1 979 (see Fig. 
UC). D, bower 19a: an arboreal tower \miIi ;.) bower perch l6'lciTi above ground. April 1979. The few 
irregularly-placed sticks to the right enc| pf the bower pwh subsequeritly beCArne a second tower. E, bower 2a: a 
tall massif supported by STlneJfcatalsblfiim 
twin tower fltniciurei 

The 25 traditional bower sites involved a total otherseasons if their stmcttire changed notably, 
of 51 bowers. These bowers were described. Photographs were taken from the same locatig?i 
photographed and/or titeasured during April and height each lime, so that temporal changes i^i 
1979, August 1984 and Febmaty 1990; ^nd in bower shape and structure could be assessed 



BOWERS OF THE GOLDEN BOWERBIRD 



299 



accurately. These data provided us with inform- 
ation on age of bowers, changes to bower 
structure and bovver building. 

RESULTS 

BOWER SITES, THEIR LOCATION AND 
DISPERSION. Traditional Bower Sites. We 
located 49 traditional bower sites (Table 1 ): 1 2 in 
SA 1 , 1 in SA2, 1 9 in adjacent extralimital areas 
(see Fig. 2), two near Birthday Creek Falls and 
six around Paluma Dam. Bower sites had 
medium to large forest trees, many saplings, and 
the odd tree ferns above and around them. Woody 
lianas and climbing pandans were common, but 
large stands of Calamus were not close to 
bowers. Canopy foliage cover was estimated 
above 37 sites: 1 2 bavins a coverage of >90%. 7 
of 80-90%, 11 of 70-80%, five of 60-70% and 2 
of 50-60%. Thus, 51% of sites had a cover of 
>80%, and81%of>70%. 

The direction of ground slope down and away 
from the bower was recorded for 45 bower sites: 
20 were on a N to E bearing of 0-90°, 1 1 on E to S 
bearing of 100-175°, 2 on a S to W bearing of 
2 1 0-240°, 8 on a W to N bearing of 280-360° and 
4 were on flatlish groimd with no slope. The 
degree of slope on which bowers were placed at 
45 sites were as follows: 1 7 on flat to 1 0° gentle 
slopes, 9 on 11 -20° slopes, 6 on 2 1 -30° slopes, 7 
on 31-40° slopes and 6 on 41-45° slopes. Thus, 
bowers were built on slopes that averaged 21 ± 
15°, with 71% being on slopes of <31°. On 0-10° 
slopes bower size (= volume; see Methods) 
averaged 740 ± 214cm^ on 11-20° slopes 589 
± 208cm^ 21-30° slopes 625 ± 223cm^31-40° 
slopes 620 ± 240cm-% and 41-45° slopes 684 ± 
393cm\ There was no significant correlation be- 
tween bower size and degree of slope buih upon 
(rs = 0.22,P>0.05). 

Bower sites were on flatter terrain and along 
ridge slopes either side of tracks or road, on 
gentle slopes and ridges immediately around the 
hill crest, and below steeper slopes where terrain 
levelled (Fig. 2). The 12 traditional sites in SAl 
averaged one per 4.2ha, and were spatially 
dispersed throushout suitable topographic types 
(CD = 1 .45, t = r.54, P >0. 1 ). Eight of the 1 2 were 
located on flat to gently sloping (<10°) ground 
(mean ^ one per 3.8ha) of topographical types 4, 
5, 6 and 8 (Fig. 3). The remaining four sites (1, 3, 
6 and 1 7) were located in topographical types 1 
and 2 (one per 3.0ha), on steeper ground. No 
bower sites were found in topographic types 3 
and 7. 



In S78, the mean inter-bower linear distance 
from one site to the next closest one in SA 1 (n = 
12), and three additional sites (7, 21 and 29) just 
outside its perimeter (see Fig. 3 ), was 1 65 ± 4 1 m 
(range 1 10-222m). Mean NND distance was 1 5 1 
± 44m (range 110-222m). Durina seasons 
S78-S90, the mean NND of bowers at these 15 
sites varied from 138 ± 52 to 151 ± 47m (mean 
of mean = 147m). Differences were due to 
temporary disuse of bower site 20 during S87 and 
S88, and the establishment of replacement 
bowers at difllerent locations within a site. 

Rudimentary Bower Sites. We found 1 1 
Ridimcntary bower sites (Table 1): ten in SAl 
(including one just outside it; see Fig. 3) and 
another 140m outside SAl. Eight were on flat to 
gentle (<n°) slopes and the others on 21-30° 
slopes. Canopy cover was 70-85% (n = 3). 
Rudimentar>' sites in SAl averaged 78 ± 36m (n 
= 8) from a traditional bower site. Of 11 
rudimentary sites, three were attended for one 
season, one for two, and foui" non-consecutively 
for two, three (n = 2) and four seasons. Three 
others were abandoned when found (pre-S78). 

BOWER CHARACTERISTICS. Structure and 
Size of Traditional Bowers. Traditional bowers 
were single or twin tower structures of sticks of 
varying lengths, texture and diameter (Fig. 1). 
Only one (the main) bower structure at any bower 
site was attended consistently, maintained and 
decorated during a season. We examined a total 
of 86 traditional bowers at 49 traditional sites: 12 
were disused old bowers ( 1 1 being twin towered) 
that had been active before S78, but the other 74 
were attended during some part of the study 
(Table 1). Of the 86 bowers, 36% had a single 
tower and 64% two towers when found. At least 
14 single tower bowers subsequently became 
twin tower bowers (see Table 4), thus increasing 
the percentage of twin tower bowers to 80. 

Single tower bowers varied greatly in shape 
and size from: a single compact conical structure 
with a cur\'ed vine bower perch (Fig. 4A); a 
single bulky, irregular-shaped large massif with a 
sloping living sapling bower perch (Fig. 4B); a 
huge amorphous muUi-peaked massif with a base 
circumference 615cm and a rotten ground bower 
perch (Fig. 4C); an entirely arboreal structure 
with bower perch 1 6 1 cm above ground (Fig. 4D); 
to a 205cm tall single massif with a Miane' bower 
perch 116cm above ground (Fig. 4E). Single 
tower bowers had only one platfonri, even though 
a handllil of sticks was placed at the opposite end 
of the bower perch in a few cases (see Fig. 4B,D) 



300 



MhMOlRS OF THE QUEENSLAND MUSEUM 




FIG Shapes niiU of twin tower bowers. A, bower :9b. a bower whh'onBt<SWer terrestrial and llie other 
aiboieal. Xiigiisi 1 484. Note: the sticks on the two bower perch platforms are conspicuously Jijor^ neatly and 
iighils aligned. B. bower 4a: a bower with one terrestrial tower much larger than ttie other. April 1979. This 
bower ctmnged little during the entire study. C\ bower 20a: a 'cUssic' compact U-shaped tMi^iftt structuEp 
wilhtwowdl-formed platforms. April 1974. Note: the sticks of theplatformstTieetort^he^jow©^ 
of the towers have met and are fused beneath it. D, bower 17a: a bower with iwo xvidety'-spaiEed but compact tall 
tovvers of similar hei«;hl built onalarp woody vine that also panly supported the towei-ontherigH^ 1979. 
E, bower 1 Oa: a bow^r witlitwo w}de^-s;>acB^amorpho?J3 towers. The vine b<jwer perch also partly supports the 
tower on the ri^t. April 397^.Thi5J>ower ^gbd IFftle during the entire study (see Table 4). Note ir is ea^v to 
see how abovfi^liJcelhfeinayhavfebrijgfnatedft^ two low arboreal subsidiaries such as the ones in Fis. 8B. F, 
bower 16^; fliSk6leiarbcfWi6^mthart>te^^ that, despite its appearance, was regularly attended troin 

878-S85 before being replaced by a new dne. April i 979. The terrestrial subsidiary in the centre background 
rtrtttdaway. 



at w hich site the second tower could be built to A few either arboreal (Fig. 5A) terrestrial 
develops Iwntawer structure. (Fig.. 5?) twin feoWetl ha*4 markedly 



BOWERS OF THE GOLDEN BOWERBIRD 



301 



TABLE 2. Size (cm) and volume (cm^) of single and twin tower traditional bowers of male Golden Bowerbirds 
on the Paluma Range, north Queensland. * - sample sizes vary because not all parameters of each bower were 
measured of each bower; ** = height (base to top apex) x base circumference. 





Single tower bowers 


Twin tower bowers 


Both towers 


Larger tower 


Smaller tower 




Height 


Base 
circum- 
ference 


Volume 
** 


Height 


Base 
circum- 
ference 


Volume 
+* 


Height 


Base 
circum- 
ference 


VoluiTie 

** 


Both 
towers 
volume 
** com- 
bined 


Distance 
between 
tower 
apices 


Shortest 
distance 
between 
tower 
bases 


Sample 
number * 


15 


13 


13 


60 


39 


39 


60 


39 


39 


39 


34 


38 


Mean 


148.6 


439.0 


646.6 


130.3 


347.1 


456.6 


82.8 


209.5 


197.0 


654.0 


98.4 


24.6 


s.d. 


32.3 


102.2 


215 


27.92 


98.4 


178.0 


44.3 


91.6 


157.6 


262.1 


20.9 


17.7 


Range 


100-205 


274-615 


320-1010 


70-190 


200-534 


186-982 


13-193 


65-390 


104-579 


309-1449 


70-150 


0-80 



asymmetrical towers. Eleven bowers had one 
larger terrestrial tower and a smaller arboreal one 
when first found. At least six arboreal towers 
subsequently became ten^estrial due to an accum- 
ulation of dropped sticks beneath the structure. 
Typically, however, twin bowers had two 
well-fonned towers with one tower taller and/or 
more massive than the other and their towers 
extending down to the ground. They varied 
greatly in shape and size: from a compact 
U-shaped structure (Fig. 5C) to widely-spaced 
neat (Fig. 5D), more amorphous (Fig. 5E), or 
skeletal (Fig. 5F) structures. The towers of the 
largest terrestrial bower were 1 74 and 1 98cm tall, 
with base circumferences of 5m and 3m 
respectively (Fig. 6A). hi two bowers both towers 
were arboreal, with their bower perches 2m 
above ground (Fig. 6B). 

Mean bower measurements for 15 single, and 
60 twin, tower bowers are given in Table 2, where 
ranges exhibit the cumulative variation of bower 
structure outlined above. Single tower bowers 
averaged 1 3% taller, 2 1 % larger around the base, 
and 29% bulkier (volume), than the larger tower 
of a twin bower. Moreover, their mean volume 
was similar to that of the mean combined volume 
of both towers of twin structures (= 654cm3). In 
5 5 twin bowers, the larger tower was both bulkier 
and taller than the smaller one, but in five bowers 
the bulkier tower was the same height (n = 3) or 
slightly shorter (n = 2; by 19 and 34cm). 
Distances between tower apices averaged 98 ± 
21cm (n = 34), and between their bases 25 ± 
18cm (n = 38). The bases of eight twin tower 
bowers were connected beneath the bower perch 
by the amalgamation of sticks of each tower (see 
Figs 5C, lOF). 



Towers were built around, and supported by, 
saplings and vines (<25cm gabh) and/or trees 
(>25cm gabh). Larger single towers, and those of 
twin bowers, encompassed more such supports 
than did the smaller tower of a twin (Table 3). The 
gabh of saplim^s within bowers averaged 7.9 ± 
5.7cm, and of trees 62.8 ± 23.6cm. Of 272 
examined tower supports, 83% were saplings, 
12% trees and 5% vines (12 woody vines and a 
Calamus vine). Four of the vines and two of the 
saplings also formed the bower perch. 

The bower perch protruded from a single tower 
bower, or connected the two towers of a twin (see 
Figs 4-6). The axis of the bower perch was at right 
angles to the axis of the inter-tower bower 
'avenue'. Bower perch compass alignment was 
recorded at 49 traditional bowers: 20 were 
aligned between 0-45^ 8 between 45-90°, 9 
between 90-135^ and 12 between 135-180^ The 
bower perch in 61 bowers consisted of: a woody 
vine (43%), living saplings leaning toward the 
horizontal (24%), a rotting dead branch or vine 
(24%) or a narrow tree root (3%; see Figs 4 & 5). 
Bower perches averaged 4.6 ± 6.5cm in 
diameter (Table 3). The top of bower perches 
averaged 42 ± 40cm (n = 59) above ground; but 
if the 4 atypically arboreal towers, with resultant 
imusually high perches, are excluded (see above) 
the average becomes 33 ± 19cm. 

Where tower sticks met the bower perch they 
were conspicuously more neatly and tightly 
aligned into what we term a platfonn(s); see Fig. 
1. Wliereas single tower bowers had only one 
platform, twin structures had a platform at either 
end of the exposed bower perch. The mean length 
of exposed bower perch of twin tower bowers 
was 1 8 ± 8cm (n = 49), but the platfomis of 4 
such bowers actually met atop the bower perch 



302 MEMOIRS OF THE QUbENSLAND MUSLUM 




FJG.6. (digest and hi^ieSttmntowtrikiw^iSi A^t^^w^ the 
l^eRst terrestri^ Win strUctuite h&d T74 stnd iv^ta fsJt tmvcrs 
with base circumferences of 5m and 3m r^epfiv^te ^ 
hon/ofital living sapling formed its bower perch. Aprft 1979. B, 
bower 34b: the highest arboreal twin siructurc had both towers 
and its vine bower perch 2ni above ground. October U395.Mote; 
the sticks of the nvo platforms eKtend to uieetand fuse atop ihe 



(sec l ijis 5C & 6B). BdfWBT decorations were 

placed only on platform slicks and llioso Jusi 
bcside/aho\e ihcni (Fig. 1 ). The reluliv e Linuniiiies 
of each decoration type and iheir placement (i.e. 
OJrt ot>e orbpthillalfonm) varied from bower to 

season, decorations cons^Bfi pSTaeatpet (30-40 

pieces) oi greyish-green Ifchen IhveO sp, and 
5-20 sprigs of creamy-while dt-hihccd ripe fruit 
\\ilh attached black seed (seed pods hereafter) of 
melicope, Melicope hroadhentium (see Fig. 1 ). 
ptliexi jess frequently used, decorations included 
Trearhy-wKte flowers of jasmine (Jasminium 

kcijcu .skii). Brov\ n Silkv Oak ( Darlin^a 
Liurlingkma) and Dcndrobium spp. orchids. 
Strmiure and Size of Rudimentary Bowers. We 
examined 1 2 rudimentary bowers at 1 1 rudiment- 
ary sites (one site had two bowers built/attended 
during diircrcni seasons; see Tuhlc i ). These 
were poorly constaictcd, maintiiincd, decorated 
and attended nrcgiilarly for iMiIy a few du}'s/ 
weekii eatJi season* by immalwe males- They 
eOBaiSte4ufa cbiiicm-sM^ Idoseitiasstif strcks 
ladtUl^-ajpbillbnn, or witB-pjO^ an ill-detmed 
onis, and often lacking a bower perch or 
decorations (Fig. 7A). Height of their towers 
averaged 84 ± 12cm (n = 9; ? being loo old u> 
measure), and meatr volume 304 j: 188cm^ (n = 
3), Only one rudimentary bower had a second 
tDWcr, a merc 50cm tall pile of sticks placed on 
the ground. 



Fbllwing the disappearsnceof their long-term 
tradition^ adult male owners, a lew tnidiiional 
sites were irrcguliirly aiiended by immature 
male(s) who built new rudimentary bower 
structures there. Ruditaeitlatj bowersat tradiiioit* 
al sites were1?etter formed, wilfe a boWfcr perolt 
and inrr: vhi pl iifrvni (see Fig. 7B), lhan 
tliosc huili ,u f iiiinii.ni iry sites. They were all 
conical single towers (mean height = 95 ± 25cm, 
n - 5: mean volume 3 1 2 ± 1 02cm, n = 2J, Some 
became a larger, ami a traditional, bower once^ 
adult rnaieilgain regularly attended the site. 

S^rwcfi^rfi j!in4 SSz^ of Suh&idmy $awer 
^tctu'r&s. Theireqiicifitiiae, by males, of one 6t 

more fa\ (Hired horizontal perches around bower 
sites resulted m bn*ds placing, or leav ing, sticks 
on them at the point lhe\ di\ i i li j-J l! (>m ihe trunk. 

Suclvsticks accucnulated, becmne fused by fungi^ 
and ihiis deVctet3rea;i <iwie, subsidtiry 
slructiires. Some subsidiaries were in ihe 
immediate vicinity of the main bower(s); (see 
Figs 4A, 5F)» "V^Ue others^ were up to 20m 
distant. 

We recorded 36 teiTcstriai and 115 arboreal 
subsidiarA structures at 46 traditit^nal bowers: 
located at an a\erage of 5.3 ± 4.2m from the 
main bower perch, and averaging 3.3 (range 
1 - 1 6) in number per site (Table 3). All but four of 
the 46 bowers had several arboreal subsidiaiy 
Structures tmean=2.7, range- 1-^13), hutpniy IS 



J30VV ERS Of THE GOLDKN BOWHRBIRD 



303 



TABl-E 3. Numbej- and meaburcnients (cm) ofbower perches, lower supports and associated subsidiaiy "bower 
sample sfaes vary because HtitaLU parfltfieters of each fewer TWtJt^tiieteuied ofirteedia.. 



Tower supports 







riphngs'vinefi. 


TreeH 






Si agio 
tosher 

hnwei-ri it 
larger 

tower of 
Iwin 
bowers 


Smaller 

Uvin 
bowers 


beight 


fowei" 
hovN'crs 1ft 

Iwin 
buwcrs 


Smaller 
twm 






Hi;il-:-.i 


L luJing 
:irhu'"iLal 
bowt'i's) 




Sample 
number * 


50 


35 


5i> 50 


35 


27 


49 


59 


55 


51 


Mean 




I--J 


T.y** i 0.54 


0,1 


62,8 


17.7 


JI.7 




4.6 




2.6 


1.3 


5.7 


,0.7 


4 




S_I 




!S K 


6o 




H2 


0-5 


M3 


0-3 


D-I 


38.5^1 


0-3 K 


3-2ijn 




I_^-1S 



hBdteitefitrial6hes(mean = 2. range= 1-5). The 
height bf-arbpreal subsidiaries, from their base to 
apex, averaged 168citi (range - 36-3S8cmj iind 
that of the terrestrial ones 69ctn H^W^ge^ - 

I5-I06em) from ground to apex. 

Terrestrial subsidiary structures were built 
mostly around SapHngs, but a few incorporated a 
tre^ (Fig.SA). Two subsidiaries wereoft^ buiU 
fiili^setcrgefteTf<lm apart) ontheSan*ehwi'zriiikI 
plinc and resembled a miniature twin tcwer 
bower (Fig. SB). A few subsidiaiy structures. 
particuUirly such pair-, i.-f ilieiii. subsequent!} 
became tlie basis for new niaiu bowers. Arboreal 
subsidiaries were built where a branch tbrked 
horizdptally from a sapling/iree trunk (Fig. 8C), 
or where a leaning sapling or vine crossed a 
sapling/tree trunk (Fig. 8D). Most (n = 00) w ere 
too high (>1.5m above ground at base) to 
measure. Those nearer the ground (<1 .5m ) were 
usually larger and conical, and averaged 61cin 
(range 20- 120cm, n = 25) tall (Fig. 8D). As 
dropped sticks accumulated beneatfi them (n - 3), 
such arboreal structures became terrestrial ones. 
We recorded only one terrestrial and three 
arboreal subsidiaries at rudimcnian, bower sites. 

Bower Sire Consumcy and Bow er A^j,c. Ol the 25 
traditional bower sites moniiored seasonally 
(S73-S97.), :84%.>V?T9 attended every season for 
20 y^ars (F}g,'2; 'Kfele 4). Of the 4 remaining 
sites: site 20 was attended for 18 (unattended 
S87-S88), site 27 also for 18 (unknown in S96, 
and derelict in 897), site 1 6 for i 1 , atJdsiteJJ ft>r 
5 (S78-S82) consecutive seasons, 

Twin i0V^| feowef s were tist^bliSshed and 
attended fbtttfe first S and 14 seasons at boMer 
sites 20 27 respectively, but when their 
Iptig-'t^tm :a4ult male joAVtreis dtsdpp^&i'eiJ 



Jemeslrial 



I A Jboregl 



2 6SS*« 



17 



H^iBtit 
.above 
eroynd 



I 

1-5 



26 



2.6 



Tol.ll 
number 
per 

site 

46 



llisiajice 
IVorn 
hnvver 
perch 



3J 
h»6 



4I_5__ 



iimmrtUTcUialcs took o\ cr and built and attended 
rudimentary bowers (2 at site 20 and I ai 27, 
Table 4). Site lb had 2 single tower traditional 
bowers attended consecutively by 2 aduh males 
pve^: U seasons (until SS8) was then 
abandt3>red (Table 4^1. There was bnly a 
rudimentary bow er at site 21 ih 578, but I4ni 
away was a di^iused large twin lower flattened by 
a tree 6! tit ftedBbviously been attended diiririg 
pi^vipus seasons. Its rudimentary bower did not 
change (S78-S82), was not replaced, and was 
only irrci:ulary attended b\ one io several 
imniaiure males bethre being ab:indoned. 

The 2^ Uadiiioital ijLes had a total pf 5 J bowers 
acthetti durbifi;tllre3md3r('Fable 4). Seve»t3f1}^i( 

25 sites had one main bower, 12 two bowers, 4 
three and 2 l our hinveTs; at a mean of 2 per bow er 
site o\cr onie. Two bowers (15a and 20d) re- 
mained ruclimeniary Igr manj; sea^pns before 
becoming larger and tra^floiral CHies{S^befC)W), 
whereas three other rudimentary bowers (2()c. 
21a, 27b} did not progress and were abandoned 
(lfaWe4). Thu5^ of the 51 bowers, 48^iv^ 
became traditional ami 3 remained rudimentary'. 
The mean tninunum hfc' of a Iradilioiiai bower 

■was 9.6 ± 6.3 (n - 4&) years. Six- trafditipnal 



304 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 7. Shapes and sizes of rudimentary bowers. A, this newly-built rudimentary bower (30) was first found at a 
new rudimentar) site in August 1979. B, this rudinientarx- bower (21a) vvas built at a traditional site after a tree 
had destroyed the previous tw in tower structure. It never increased in size and the site was eventaully abandoned. 



bowers were attended for a minimum of 20 seasons 
(Table 4). 

BOWER BUILDING AND STRUCTURAL 
CHANGES. We observ ed construction of 1 trad- 
itional bowers. Four ( 1 a, 1 7b, 45b, 45c) started as 
completely new arboreal strucltires, 4 (3b, 6b, 
20b, 47b) developed from an existing subsidiary 
bower structure, and 2 ( 1 5a, 20d) were built trom 
existing rudimentary bowers, as follows: 

In S78 we fotind only a derelict (pre-S78) 
bower at site 1 . In March 1 979 a handful of slicks 
had been newly-placed on a fallen horizontal 
sapling where it met a vertical tree, 20m distant 
from the derelict structure. This new bower (la) 
resembled a sparse arboreal subsidiary, but it was 
decorated with two sprigs of melicope seed pods 
and one piece of lichen placed on Freycinetia sp. 
vine above the bower perch (Fig. 9A). By April 
1980 the structure vvas a small conical arboreal 
subsidiary. By June 1980 it was an untidy tower 
of unfused sticks, lacking a platform and 
resembling a terrestrial rudimentary bower 
decorated with 10-12 lichen pieces (Fig. 9B, 
compare with Fig. 8A). By S80 it was a small 
terrestrial single tower, and by S8 1 a larger massif 
with a well-formed platform. Its bovver perch 
subsequently slipped to the ground and a small 
handful of sticks placed on it 1 4cm from the large 
tower. By S82 this bower w as a terrestrial twin, its 
second tower much the smaller (Table 4). It took 
3.5 years to reach this stage. 

Bower 17a was flattened by a tree fall in S85. In 
November 1986 we found a small conical 
tirboreal bower ( 1 7b), of loosly-placed unfused 
sticks, where a leaning sapling crossed a vertical 
one, about 45m from the flattened bovver and 
63cm above ground. By October 1987 this was a 



terrestrial single tower, and a pile of sticks had 
been placed fiirther along (20cm) the sapling 
where it crossed a large tree. By February 1990 
the latter pile of sticks was a small second 
arboreal tower, and by November 1991 this was 
larger and terrestrial, h thus took four years for 
this bower to become a large twin tower structiare 
(Table 4). 

In December 1979 we found a newiy- 
conslructed small single conical arboreal tower 
of unfijsed sticks, piled between the vertical 
trunks of three saplings, 50cm above ground. 
This new bower (45b) was 20m from a derelict 
one of the previous season (Table 4). In S80 
bower 45b was still small, but by S81 was a 
substantial terrestrial single tower bower that 
remained so until S88, but by which time it had 
dclcriorated. In October 1 989 replacement bower 
45c, about 20m from 45b, was a single tall tirboreal 
tower 60cm above ground. By September 1 990 it 
was a twin tower bower with its second tower an 
arboreal one. By November 1991 both towers 
had become terrestrial (Table 4). 

Four traditional bowers developed from an 
existing arboreal (6b and 47b) or terrestrial (3b 
and 20b) subsidiary bower (Fig. 9C,D). These 
took two to four years to become large single 
(47b) or twin (3b, 6b, 20b) tower structures (see 
Table 4). Traditional bower site 1 5 was attended 
only by immatures males from S78-S87, and its 
bower was small and rudimentary. When an adult 
male took over the site in S88 the rudimentary 
structure, unchanged for many years, became a 
larger single tower bower. By S90 it was a twin 
structure (see Table 4). Similarly, in S89 bovver 
2()d was a new loosely-constructed rudimentary 
terrestrial tower attended by immature males. It 



BOWERS OF THE GOLDEN BOWERBIRD 



305 




riG 8. Shapes and sii^es i>rsubsit]ian bovver structures. A, this it-rrestrial subsidiary vvas4in tVom the ftiuin bower 
slruciurc (2()a) in April 1979. When bower ZOa (sec Fig. 5C) was damaged bv n ncc fall in January 1981, Ihis 
terrestrial slaiclure formed the basis ofrhc new bower (20b). By SX2 it wa^ a uvin lower bower. B. oflen (wo 
arboreal subsidiaries \vcre buiil close logclher ( I m apart) on the same horizontal support so that xlie> 
Eeserabled a miniature twin tower bower. AVil 1 '^79. C , arboreal subsidiaries v\ erc usually built w here a t)rancli 
forked horizontally from the trunk of a saplingtree, April 1 979. diis arboreal subsidiary was huili where a 
viiiB crossed thetruiAofa smaJl tree. April ] 979. Note: This subsidiary deteriorated but could have become an 
aAoreaJ bower bUeIi as the me shown in Fig. 4D. 



remained so until at least S93, but by S97 ii was a 
substantial single tower bower (its ownership 

uncon tinned). 

]^amerpus bowers progressed through sts^es 
of stntcturallieveloptrtent sitoiilarixJ the above. 

Of the 48 traditional bowers examined at 25 
traditional sites, 20 were initially single lowers, 
but 14 of them were changed subscquetitly intQ 
twin structures (Table 4 and Fig. lOA-D). The 
original lower of these bffWer^ teitiaSned fhe 
larger oflhe Iw^^. Morcoven the main towers of 8 
arboreal (2 single and 6 twin) bowers 
subsequently became tentstliai on«s» as did the 



smaller lowers of 6 twin tower bowers (Table 4 
and Ffe 10B,F). 

Must second lowers sinned as arboreal 
structures, because the bower pereJt was above 
grtiurtd. For example^ bower I9a was an arboreal 
(128cm tall) single tower in S7S wilh but a 
handful of sticks at the far end of its huwcr perch 
(see Fig. 4D). Heavy rains in January l9tSl 
caused the entire bower structure to slip toyvijrtj 
the ground (bower perch frdra 161cm dowii to 
60cm). In SHZ it was an arboreal twin b(nvcr, the 
handful of slicks having been developed into the 
second tower, but both towers soon became 



306 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 9. Bower construclion at traditional sites. A,B, bower I a: this was little more than a handful of newly-placed 
sticks on a fallen horizontal sapling where it met a vertical tree in March 1979 (A) but by June 1 980 (B) tliis had 
become a terrestrial tower of unfused sticks that lacked a platfonn. By September 1982 it was a Uvin lower 
structure. C, bower 6b: this originated in March 1980 from an arboreal subsidiar>'. The subsidiary formed the 
main tower and a second smaller tower was subsequently built al the opposite end of the bower perch about a 
small ti-ee (right). This photograph was taken in August 1984, three and half years since building was 
commenced. D, bower 3b: this originated from a terrestrial bulky may pole- shaped subsidiaiy May 1982. The 
subsidian* became the main tower and the handful of sticks on the 'right hand end of the bower perch later 
became the second tower/platform. 



terrestrial, as dropped sticks accumulated 
beneath them. Similarly, bower 2a obviously 
started as an arboreal single tower, but sticks 
steadily accumulated beneath it until they 
reached its base and thus formed a huge single 
terrestrial massif (see Fig. 4E). It became even 
larger during subsequent seasons, but it was not 
until S90 that we noted the beginnings of a second 
tower at the opposite end of its bower perch (see 
Table 4). 

The height of some single, and the larger of 
some twin, towers changed surprisingly little 
from season to season (Fig. II A, B), whereas 
others increased in bulk as slicks were added 
(Figs ] 1C,D). Some reached the same or greater 
height (but not the bulk) of the larger tower^^Table 
4). After several seasons, some bowers 
deteriorated and became smaller as their towers 



decomposed or collapsed (Table 4). For example, 
in S78 bower 27a was a 'classic' twin tower 
structure with w^ell-fonned platforms (Fig. 1 IE). 
By February 1990 it had deteriorated, and its 
bower perch collapsed (Fig. 1 IF). It w^as replaced 
in S92, as were bowers 19a and 22b, after the 
extremely dry S9 1 . Replacement was due to gen- 
eral bower deterioration, including the collapse 
of a main tow er support and/or bower perch (n = 
13), tree fall (n = 6) or mammal damage (n = 2). 
The mean distance of a replacement bow er from 
the replaced one was 14.3 ± 12.7m. 

We did not seasonally monitor subsidiar}' 
bower structures, but in August 1984 w^e did note 
that most of those recorded in S79 were 
deteriorating, or had disappeared, and new ones 
had replaced Ihem at other locations about the 
main bower on the bower site. 



BOWERS OF THE GOLDEN BOWERBIRD 



307 



DISCUSSION 

We discuss our results in the light of previous 
Golden Bowerbird bower studies, and compare 
them mostly with data for the closely related 
rainforest-dwelling bowerbirds Scenopoeeies, 
Archboldiam&Amblvomis spp. (see Kusmierski 
etal, 1993). 

BOWER SITES, THEIR LOCATION AND 
DISPERSION. Traditional bower sites of Golden 
Bowerbirds were found on flattish to gently 
sloping ground along ridge tlanks above steeper 
slopes, and mostly with >70% canopy cover 
above. None occurred on hill or ridge crests, or in 
disturbed forest dominated by Ccdamus palms 
and creek lines and their adjacent, typically 
steeper, slopes were avoided. Bowers have been 
described as occun*ing in similar sites on the 
Atherton Tableland (Day in North, 1 904; Bourke 
& Austin, 1947; Chisholm & Chaffer, 1956; 
ChatTer, 1958; Warham. 1962; GiUiard, 1969; 
Crome & Moore, 1989; Frith & Frith, unpubl. 
data). 

Bowers of the closely related gardener bower- 
birds in New Guinea occur mostly on ridge crests 
or slopes below them (Simson, 1907; Rand in 
Mayr & Rand, 1937; GiUiard, 1969; Schodde & 
McKean, 1973; Diamond, 1972, 1987, 1982a). 
Pruett-Jones & Pruett-Jones (1982) examined 46 
active bowers of Macgregor's Bowerbird, Amhly- 
amis macgregoriae, on Mt Missim. Kuper Range 
and found 87% on ridge crests and the remainder 
3-30m below crests on relatively level areas of 
the slope. They concluded that the habitat 
variables influenced the choice of bower site by 
Macgregor's Bowerbird rather than the selection 
of the ridge itself They found such things as 
degree of canopy closure (>80%), slope and 
width of the ridge important factors for site 
selection. They found 42 bowers of Macgregor's 
Bowerbird spaced linearly and regularly along 
ridge crests at an inter-bow er distance of 169 ± 
64m. This figure is comparable with our Golden 
Bowerbird linear inter-bower measurement of 
169 ± 40m,ratherthanourNNDof 151 ± 44m 
(see RESULTS). Diamond (1987) estimated that 
distances between five Vogelkop Bowerbird, 
Amblyornis inornatus, bowers were several 
100m. He pointed out that this was similar to the 
inter-bower spacing in Macgregor's Bowerbird 
and the 0,5 km separation for eight bowers of the 
Golden-fronted Bowerbird, A. Jlavifrons 
(Diamond, 1982a). 

Dispersion in Macgregor's Bowerbirds appears 
to be largely the resuh of socially interacting 



males utilizing available favoured topography. 
Its mating system has been characterized as being 
intennediate between lek behaviour and teiritor- 
iatity, with birds maintaining even dispersion in 
part by 'buffering their display space against 
intruder pressure' (Pruett-Jones & Pruett-Jones, 
1982; pers. obs.). Traditional bower sites of the 
Golden Bowerbird averaged one per 4.2ha and 
were spatially dispersed, not clumped (contra 
GiUiard 1969: 321), throughout suitable 
topography. Male Golden Bowerbirds disperse 
their bower sites over suitable topography and 
habitat in an CA'en way, similar to Macgregor's 
Bowerbird and apparently as a result of a similar 
social system. Dispersion of the bowers of 
Archbold's Bowerbird is also relatively even 
tliroughout suitable habitat (Frith et al., 1996a) 
and not clumped into leks (contra Diamond, 
1982b). 

Leks have been defined as requiring the 
following characteristics: clumped distribution 
of males; the ability of females to freely choose 
mates; no parental care by males; and no 
resources of value to females available at male 
display sites other than spenn (Bradbury, 1981). 
True lek behavioiu" has not been demonstrated in 
any bowerbird species (Donaghey, 1981 ; Pruett- 
Jones & Pruett-Jones, 1982; Diamond, 1986a; 
Borgia, 1986; Oakes, 1992; Lenz, 1993; Frith & 
Frith, 1995; Frith et al, 1996a,b, this study), as 
this requires that males at their bowers be in visual 
contact (Frith & Beehler, 1998). Rainforest- 
dwelling Tooth-billed Bowerbii ds may be the only 
exception, as courts on the Paluma Range showed a 
dispersion intermediate between an even spread 
and true (i.e. exclusive) clumping (true lek) over 
suitable habitat (Frith & Frith, 1995). It remains 
to be demonstrated conclusively, that 'clumping' 
of male Tooth-bill courts does form a lek, albeit 
an exploded one. It is possible the dispersion of 
courts was the result of males utilising the only 
appropriate topography available, as appears to 
be the case in Golden and gardener bowerbirds. 

Rudimentary' bower sites and structures were 
short-lived, built and used sporadically by one or 
more immature males during one, several 
consecutive, or non-consecutive seasons. Similar 
rudimentary stmctures have been described for 
Macgreaor's (Pruett-Jones & Pruett-Jones, 
1982), Regent (Chaffer, 1 984; Lenz, 1993), Satin 
(Vellenga, 1970; Donaghey, 1981; Borgia, 1986), 
Archbold's (Frith et al., 1996a), Great and 
Spotted (Frith & Frith, unpubl. data) Bowerbirds. 
Rudimentary bowers of Amblyornis spp., probably 
built by younger males, are often found at lower 



308 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG ] 0. Seasonal changes to the towers of traditional bowers. A, B, howcr 34a: this was a single lower bower in 
April 1979 (A) but by August 1984 (B) was a Uvin tower strueture with its main tower then 180cm tall. C,D, 
bower 8a: this was a single tower bower in April ] 979 (C) but by Februaiy 1 990 (D) was a massive twin tower 
structure. E,F bower 33a: this had one ofils towers small and arboreal in April 1 979 (E) but by August 1 984 (F) 
both towers were terrestrial. Note: sticks of both towers were fused beneath the bower perch. 



altitudes than the traditional bower structures 
presumably built/owned by older males 
(Diamond, 1 986b, 1987 and references therein). 

BOWER CHARACTERISTICS. Previously the 
bow er of the Golden Bowerbird was thought to 
consist only, or typically, of two towers, and with 
one low er usually taller than the other (De Vis in 
Meslon, 1 889; "Campbell, 1900; Meston in 
Mathews, 1926; Cooper & Forshaw, 1977; 



Johnsgard, 1994; Schodde & Tideman, 1988; 
Donaghey, 1996). Oiu* findings clearly show that 
bowers may be of one or two towers, and that 
their size and shape varies greatly (Table 4). In 
the case of traditional bow ers, the structure of a 
single tower averaged some 20% larger than the 
average size of the larger tower of a t^in strucliire. 
Moreover, its mean voltune was similar to that of 
the mean combined volume of both towers of a twin 



BOWERS OF THE GOLDEN BOWERBTRD 



309 



structure. Thus, single tower bower structures may 
demonstrate to conspecifics that the owning male 
has expended similar effort in building as have 
males constructing a twin tower bower 

Golden Bowerbird bower perches or their 
'avenues' did not exhibit a pattem of compass 
orientation, as is the case in several true avenue 
bower builders in which the avenue is aligned on 
or about the north-south axis (see Frith et al., 
1996b). This orientation apparently enJiances 
illumination of bower decorations and the dis- 
playing males (Marshall, 1954; Frith et al, 
1996b). We did fmd that almost twice as many 
bower perches of Golden Bowerbirds were 
orientated to within 45*^ of the north-south axis, 
or the north-south half of the compass, than were 
to within 45° of east-west, or the east-west half of 
the compass rose. Thus, given bower perches 
were at right angles to the avenue between twin 
tower bowers, the orientation of the ^avenue' was 
predominantly within the east-west half of the 
compass. We can offer no explanation for these 
observations at present. 

Sticks of Golden Bowerbird bowers, other than 
recently placed ones, become firmly fused 
together by the action of a fimgus (Mathews, 
1926; Chisholm & Chaffer 1956; Warham, 1 962; 
Frith 1989, this study) ubiquitous to the lower 
forest sub-canopy (Jackson in Chisholm, 1957). 
Certainly, birds do not glue sticks together with 
saliva, or anything else, as suggested by some 
authors (e.g. Schodde, 1976; Diamond, 1987; 
Schodde & Tidemann, 1988). 

Our long-term observations of rudimentary 
and traditional bowers indicated that most bower 
sticks are placed in a somewhat dishevelled 
fashion, resulting in great variation in bower 
shape and bulk. Their untidy construction sug- 
gests gross bower features are of less significance 
to females than is the discrete part of them 
modified into a 'platform(s)' for the exclusive 
placement of decorations. While traditional 
single or twin towered bowers varied greatly, 
they all had a conspicuous platfonn of more 
carefully and better aligned finer sticks to one 
end, or both ends, of the display perch. In view of 
bower structure quality in other bowerbirds 
(Borgia 1985, 1995; Borgia et al., 1985), it is 
possible that the quantity and quality of 
sticks/construction incorporated into the bower 
platform(s) is of significance to mate selection by 
females. Older and more dominant male Satin 
Bowerbirds that retain more bower decorations 
mate more often (Borgia 1985, 1995; Borgia et 
al., 1985). This suggests that bower platfomi(s) 



and their decoration represent characters of 
significance in female Golden Bowerbird mate 
selection. For a discussion and review of the 
significance of bower decoration, see Frith & 
Frith (2000a). 

Broadbent (in Mathews, 1926) noted that larger 
main bowers of Golden Bowerbirds were 
surrounded by several 'gunyahs', dwarf-like hut 
structures, that we term subsidiary structures. 
Buhner (in Gilliard, 1969: 305-6) reported similar 
subsidiary structures in Macgregor's Bowerbird. 
Of 151 subsidiary bower structures we recorded, 
76% were arboreal and the remainder may have 
been originally arboreal. Often two such 
subsidiary structures, placed at an interval along 
the same length of horizontal branch, resembled a 
diminutive bower (Fig. 8B). Four such subsidiary 
structures had sticks added to them to sub- 
sequently replace, and become, the main bower. 

It is possible that some subsidiary structures, 
around the main bower, at a traditional site are 
'the casual products of social activity in 
non-breeding periods' (Chisholm & Chaffer, 
1956: 13). Sharp (in Chisholm, 1929) claimed 
that only (adult) male Golden Bowerbirds 
attended large bowers and that subsidiaiy struc- 
tures are built by females, but this is eiToneous 
and may be a result of misidentification of 
female-plumaged, immature, males at such 
structures. It is our experience that these are 
initiated by the traditional bower owner, as a 
result of a bird leaving sficks at a favoured 
singing/perching perch(es). Adult males actively 
decorated only their single main bower structure, 
but would occasionally temporarily leave the odd 
decoration on a subsidiary one. 

BOWER SITE CONSTANCY, BOWER AGE, 
BOWTR BUILDING AND STRUCTURAL 
CHANGES. Most (84%) Golden Bowerbird 
bower sites were attended over 20 consecutive 
courtship seasons, predominantly by adult males 
(Table 4 and Frith & Frith, unpubl. data). Bower 
sites of Satins have persisted for up to 30 years 
(Vellenga, 1980), Spotteds for 13 years (Frith & 
Frith, unpubl. data), Greats for 13 years (Frith et 
al., 1996b), Tooth-bills for 20 years (Frith & 
Frith, 1995; unpubl. data) and Archbold's 
Bowerbird for 11 years (Frith et al., 1996a). 

The mean minimimi active iife' of a Golden 
Bow erbird traditional bower structure was 9.6 ± 
6.3 (n - 48) years. The main causes of structure 
replacement were deterioration due to age, the 
collapse of a tower(s) resulting from loss of 
supporting plants, or a falling tree directly 



310 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1 1 . Seasonal clianges to the shape oftraditional twin lower bowers. A,B, bower 3a: this changed little in size 
and shape beUveen April 1979 (A) and February 1990 (B) despite the coliapse of the dead trunk that was 
suppoiling the main lower. C,D, bower 22b: this changed little in overall shape from between September 1 979 
(C) and Februan- 1 990 (D), save becoming more massive. E,F, bowser 27a: this changed dramatically in size and 
shape between April 1979 (E) and Febriiar>' !990 (F), becoming smaller as its towers decomposed and 
collapsed. 



damaging the bower and'or opening the canopy 
above the bovver site. The larger traditional slick 
maypole bowers of Amblyornls spp. also persist 
year to year (Pruelt-Jones & Pruett-Jones, 1982, 
1983; pers. obs.). Conversely, bowers of the 
avenue-building Chlamydera, Sericiihis and 
Ptilonorhynchiis bowerbirds are refurbished and 
reused, or are replaced annually at the traditional 



bower site, but not always at the same location 
(Velienga, 1980; Donaghey, 1981: Lenz, 1993, 
and references therein). Male Tooth-billed 
Bowerbirds annually re-create their court, more 
often than not in exactly the same place (Frith & 
Frith, 1994, 1995). Archbold's Bowerbirds 
typically renovate bowers at the beginning of 
each season (Frith et al., 1996a). 



BOWERS OF THE GOLDEN BOWERBIRD 



311 



Earlier descriptions of Golden Bowerbird 
bower building were simplistic. For example. 
Day (in North, 1904) and Marshall (1954) noted 
that when a bower is first built it consists of sticks 
and twigs placed around two small trees growing 
about a metre apart. Each season sticks are added 
lo the structures around two saplings until they 
are joined to fomi a U-shaped structure. In the 
centre, near the bottom of the U, a horizontal 
vine, stick or root is left bare (i.e. the bower 
perch). Our findings clearly demonstrate, how- 
ever, that males in fact started each bower 
structure at a point above ground, where a leaning 
or horizontal branch (sapling, vine, fallen 
branch) crosses a vertical sapling or small tree 
trunk. New main bower structures started as 
small, single, arboreal, conical or maypole-shaped 
structures. They typically became fully terrestrial 
later, when dropped/fallen sticks accumulating 
on the forest lloor reached their bases. Some 
bowers remained single tower bowers while 
others continued to grow into twin towers taking 
two to three seasons for them to reach full size. 

Tow ers of some traditional bowers changed in 
shape and size from one season to the next, and 
often incorporated more saplings as they 
increased in size, whereas towers of others 
changed little from one season to the next. 
Dropped sticks may accumulate beneath the 
bower perch to there fuse to form a solid w all or ^a 
sort of hedge^ (cf Chisholm & Chaffer, 1956: 
1 1); thus reinforcing the false impression of the 
structure originating on the ground. That most, if 
not all, Golden Bowerbird bowers originated 
above groimd is a significant finding, given that 
all bowerbird species were thought to begin 
bower construction on the groimd until Borgia & 
Sejkora (in Kusmierski et al, 1997: 310) stated 
that the Vogelkop Bowerbird builds its bower 
'from the top down'. Thus the 'foundation' of the 
Golden (and Vogelkop?) Bowerbird's bower is 
not the clearing/cleaning of an area of ground 
(contra Stresemann 1953). 

Diczbalis (1968) noted that male Macgregor's 
Bowerbirds start to clear a space around a young 
sapling, plucking olTits leaves, bringing moss to 
form a basal ring around the sapling and 
'trimming the space between base of sapling and 
the outside ring till it is clean and level. At the 
same time, the bird was bringing in its beak dry 
sticks and arranged these with its beak into spoke 
like shape around the sapling' to form a tower. He 
noted the structure was completed within a 
month, but would be improved and strengthened 
throughout the display season. Considering the 



arboreal beginnings of Golden Bowerbird bowers, 
clarification of bower development of the closely 
related Amblyoniis species would be valuable. 
Were the original bowers of Amhlyorms and 
Archboldia spp. arboreal, or has Prionodnra 
'raised' its point of initial bower construction 
from the terrestrial forni of its ancestors? 

SIGNIFICANCE OF BOWER FORM AND 
ADULT MALE PLUMAGE IN THE GOLDEN 
BOWERBIRD. The bowers, their decorafion, the 
levels of attendance at them by males, and the 
plumage morphology' and courtship displays of 
the Golden Bowerbird are of particular interest 
within the bowerbirds with regard to the 
'transferral effect' postulated by Gilliard (1956, 
1969). This theoretical effect suggests that, 
within several bowerbird genera, the degree of 
ornate/colourful plumage in adult males is 
inversely proportional to the complexity of their 
bowers. Thus, males of species developing more 
complex bowers, as external symbols of their 
dominance/fitness, have been able to replace 
their personal, and possibly costly (in making 
them conspicuous to predators), plumage 
ostentation with a bower structure and its 
decoration. The more impressive examples of 
this relationship occur \\'\ih\nAniblyornis and the 
Senculus-Ptilonorhynchus-Chlamydera clade. 
While the Golden Bowerbird is clearly most 
closely related to, and originated from ancestral, 
gardener bowerbird stock (Schodde, 1976; 
Sibley & Monroe, 1990; Kusmierski et ah, 1993, 
1997) it does not conform to the transferral effect 
discernible within these maypole builders. The 
maypole bower of the Golden has certainly lost 
some of the intricacies of Amblyornis bowers, in 
that it lacks a terrestrial moss base 'dish' (as in 
Macgregor's and Golden-fronted Bowerbirds) 
and its sticks do not form a 'hut' roof over a moss 
'lawn' or ^court' (as in Streaked subalaris and 
Vogelkop Bowerbirds). Nevertheless it is a 
massive stick structure, with a discretely located 
platform(s), the construction of which is 
commenced above the ground. To what extent the 
arboreal point of initial bower construction is 
related to the significantly divergent adult male 
plumage in Golden Bowerbirds merits 
investigation. 

Given its bower and, for present purposes, 
considering the Golden Bowerbird a member of 
Amblyornis, the transferral effect would lead one 
to predict a drab adult male plumage; at least no 
more colourfully ornamented than are the 
yellow- and orange-crested (but otherwise dully 



312 



MEMOIRS OF THE QUEENSLAND MUSEUM 



plumaged) simple maypole-building Golden- 
fronted and Macgregor's Bowerbirds. How then 
is the, apparently contradictory, massive 
maypole-bower building yet brilliantly-plumaged 
adult male Golden Bowerbird to be interpreted? 
Its colourfiil plumage is not dorsally confined to a 
crest, as in gardener and Archbold's Bowerbirds, 
but is also extensive on the nape and tail feathers. 
Moreover, the entire ventral surface of the bird is 
brilliantly colourflil. This extensive colourfiil 
pigmentation of both dorsal and ventral plumage 
is, among bowerbirds, more reminiscent of adult 
male regent bowerbirds {Sericulus spp.). Adult 
male Regent Bowerbirds perch on exposed forest 
canopy branches, to advertise their bower 
location, and subsequently descend to the bower. 
During this initial advertisement, and descent, 
their bright plumage is doubtless conspicuous to 
females. Thus, we concur with Schodde's (1976) 
suggestion that, while bower-based courtship has 
apparently ornamented/coloured the dorsal 
plumage of more terrestrially-displaying adult 
male bowerbirds, the morphology of adult male 
Golden Bowerbirds, with bright underparts, 
reflects its elevated bower perch. It also reflects 
an extensive courtship flight display (Frith & 
Frith, 2000a). We view the bright central crown 
patch and the nape patch of the adult male Golden 
Bowerbird as homologous to the extensive crest 
of the gardener (especially Amblyornis flavifrons 
and A. macgregoriae) and Archbold's Bower- 
birds. These characters, together with the brilliant 
yellow long forked tail, entire underparts, and 
pale iris lead us to concur with Kusmierski et al. 
(1993) in considering the Golden a highly orna- 
mented bowerbii d ( contra Moller & Cuervo, 1 998). 

ACKNOWLEDGEMENTS 

The first three years of these studies were 
performed by CBF as a post-graduate student of 
Monash University, Melbourne. For this 
opportunity he is most grateftil to Alan Lill for 
support, encouragement, advice and friendship 
and to the then Zoology Department, Monash 
University for practical help. The initial three 
years were financed in part by National 
Geographic Society Grants 1709 cmd 1870 to 
Alan Lill, Monash University. Andree Griffin 
provided valued company and assistance in the 
field in various ways. We thank Derek Goodwin 
for most kindly translating Stresemann (1953) 
from the German. 



LITERATURE CITED 

BLAKERS, M., DAVIES, SJ..r.F. & REILLY, RN. 

1984. The atlas of Australian birds. (Royal 

Australasian Ornithological Union & Melbourne 

University Press: Melbourne). 
BORGIA, G. 1985. Bowers as markers of male quality. 

Test of a hypothesis. Animal Behaviour 35: 

266-271. 

1986. Sexual selection in bowerbirds. Scientific 
American 254: 70-79. 

1 995. Why do bowerbirds build bowers? American 
Scientist 83: 542-547. 
BORGIA, G, PRUETT-JONES, S. & PRUETT JONES, 
M. 1985. The evolution of bowers as markers of 
male quality. Zeitschrift Tierpsychologie 67: 
225-236. 

BOURKE, R A. & AUSTIN, A. F. 1947. The Atherton 
Tableland and its avifauna. Emu 47: 111. 

BRADBURY, J.W. 1981. The evolution of leks. Pp 
1 38-1 69. In Alexander, R.D. & Tinkle, D.W. (eds) 
Natural selection and social behavior: research 
and new theory. (Chiron Press: New York). 

BROADBENT, K. 'l902. Field-notes on the birds of 
Beilenden-Ker, Queensland. Ibis: July. 

CAMPBELL, A. L. 1900. Nests and eggs of Australian 
birds. (The author: Melboume). 

CHAFFER, N, 1958. Additional observations of the 
Golden Bower-bird. Emu 58: 133-137. 
1984. In quest of Bower Birds. (Rigby: Adelaide). 

CHISHOLM, A.H. 1929. Birds & green places. (Dent 
& Sons: London). 
1 957. Concerning the Golden Bowerbird. Emu 57: 52. 
1 963. The Golden Bower-bird al the nest and bower. 
Emu 62: 251-256. 

CHISHOLM, A.H. & CHAFFER, N. 1956. Observ- 
ations on the Golden Bower-bird. Emu 56: 1-39, 

CLARK, RJ. & EVANS, F.C. 1954. Distance to nearest 
neighbour as a measure of spatial relationships in 
popiulations. Ecology 35: 445-453. 

COOPER, WT. & FORSHAW, J.M. 1977. Birds of 
Paradise and Bower Birds. (Collins: Sydnev). 

CROME, F.H.J. & MOORE, L.A. 1989. Display site 
constancy of bowerbirds and the effects of 
logging on Mt. Windsor Tableland, North 
Queensland. Emu 89: 47-52. 

DIAMOND, J.M. 1 972. Avifauna of the Eastern High- 
lands of New Guinea. Publications of the Nuttall 
Ornithological Club No. 12: 1-438. 
1982a. Rediscovery of the Yellow-fronted Gardener 

Bowerbird. Science 216: 431-434. 
1982b. Evolution of bowerbirds' bowers: animal 

origins of aesthetic sense. Nature 297: 99-102. 
1986a. Biology of the birds of paradise and 
bowerbirds. Annual Review of Ecology and 
Systematics 17: 17-37. 
1986b. Animal art: variation in bower decorating 
style among male bowerbirds Amhlyomis 
imrnatus. Proceedings of the National Acadamy 
of Sciences USA 83: 3042-3046. 

1 987. Bower building and decoration by the bowerbird 
Amblyornis inornatus. Ethology 74: 177-204. 



BOWERS OF THE GOLDEN BOWERBIRD 3 13 



DICZBALIS, S. 1968. Observations on the Crested 
Bowerbird Amhlyornis macgregoriae. 
Miscellaneous Repod of the Yamashina Institute 
5: 199-201. 

DON AGHEY, RH. 1 98 1 . The ecology and evolution of 

bowerbird mating systems. (Unpubl. PhD thesis: 

Monash LIniversit>', Melbourne). 
1996 Bowerbirds. Pp, 138-I87. In Strahan, R. (ed). 

Finches, bowerbirds and other passerines. 

(Angus & Robertson: Sydney). 
FRITH, C. 1 989. A construction worker in a rainforest. 

Birds International 1 : 29-39. 
FRITH, C.B. & BEEHLER, B.M. 1998. The Birds of 

Paradise: Paradisaedae. (Oxford Universit>' Press: 

Oxford). 

FRITH, C.B. & FRITH, D.W. 1989. Miscellaneous 
notes on the bowerbirds Chlamydera cervimventi'is 
and C. lauterhachi (Ptilonorhynchidae) in Papua 
New Guinea. Australian Bird Watcher 13: 6-19. 

1993. Courtship display of the Tooth-billed Bower- 
bird, Scenopoeetes dentirostris; its behavioural 
and systematic significance. Emu 93: 129-136. 

1994. Courts and seasonal activities at them by male 
Tooth-billed Bowerbirds, Scenopoeetes 
dentirostris (Ptilonorhynchidae). Memoirs of the 
Queensland Museum 37: 121-145. 

1995. Court site constancy, dispersion, male survival 
and court ownership in the male Tooth-billed 
Bowerbird, Scenopoeetes dentirostris (Ptilono- 
rhynchidae). Emu 95: 84-98. 

1998. Nesting biology of the Golden Bowerbird 
Prionodura newtoniana endemic to Australian 
upland tropical rainforest. Emu 98: 245-268. 
2000a. Bower attendance and behaviour at them by 
male Golden Bowerbirds, Prionodura 
newtoniana (Ptilonorhynchidae). Memoirs of 
the Queensland Museum 45(2): (this issue). 
2000b. Home range and associated sociobiology and 
ecology of male Golden Bowerbirds Prionodura 
newtoniana (Ptilonorhynchidae). Memoirs of the 
Queensland Museum 45(2): (this issue). 

FRITH, C.B., FRITH, D.W. & MOORE, G.H. 1994. 
Home range and extra-court activity in the male 
Tooth-billed Bowerbird, Scenopoeetes 
dentirostris (Ptilonorhynchidae). Memoirs of the 
Queensland Museum 37: 147-154. 

FRITH, C.B., BORGIA, G. & FRITH, D.W. 1996a. 
Courts and courtship behaviour of Archbold's 
Bowerbird Archholdia papuensis in Papua New 
Guinea. Ibis 138:204-211. 

FRITH, C.B., FRITH, D. W. & WIENEKE, J. 1996b. 
Dispersion, size and orientation of bowers of the 
Great Bowerbird Chlamydera nuchalis 
(Ptilonorhynchidae) in Townsville City, tropical 
Queensland. Corella 20: 45-55. 

GILLIARD, E.T. 1956. Bower ornamentation versus 
plumage characters in Bower-birds. Auk 73: 
450-451. 

1969. Birds of paradise and bower birds. 
(Weidenfeld & Nicolson: London), 



JACKSON, S.W. 1909. In the Barron River Valley, 
North Queensland. Emu 8: 233-285. 

JOHNSGARD, PA. 1994. Arena birds. (Smithsonian 
Institution Press: Washington). 

KUSMIERSKI, R, BORGIA, G., CROZIER, R. & 
CHAN, B, 1993. Molecular information on 
bowerbird phylogeny and the evolution of 
exaggerated male character. Journal of 
Evolutionaiy Biology 6: 737-752. 

KUSMIERSKI, R., BORGIA, G., UY, A., & 
CROZIER, R.H. 1997. Labile evolution of 
display traits in bowerbirds indicates reduced 
effects of phylogentic constraint. Proceedings of 
the Royal Society of London B 264: 307-3 1 3. 

LENZ, N.H.G. 1993. Behavioural and reproductive 
biology of the Regent Bowerbird Sericulus 
chr\'Socephaius (Lewin, 1808). (Unpubl. PhD 
thesis, Griffith University, Brisbane). 

MARSHALL, A.J. 1954. Bower-birds — their displays 
and breeding cvcles. (Clarendon: Oxford). 

MAYR, E. & RAND, A.L. 1937. Birds of the 
1933-1934 Papuan expedition. Bulletin of the 
American Museum ofNatiu"al History 73: 1-248. 

MATHEWS, G.M. 1926. Birds of Austi-alia. Vol. 12. 
(Witherby; London). 

MESTON, A. 1889. Bellenden-Ker Range (Wooroo- 
nooran). North Queensland. Parliamentary Paper: 
Brisbane. 

M0LLER. A.R & CUERVO, J.J. 1998. Speciation and 
feather ornamentation in birds. Evolution 52: 
859-869. 

NIX, H.A. & SWITZER, M.A. 1991. Rainforest 

animals — atlas of vertebrates endemic to 

Australia's wet tropics. Kowari I: 1-112. 
NORTH, A. J. 1904. Nests and eggs of birds found 

breeding in Australia and Tasmania. Vol. 1. 

(Australian Museum: Sydney). 
1909. Notes on Newton's Bowerbird. Februray. 

Victorian Naturalist 25: 160-1 68. 
1914. Nests and eggs of birds found breeding in 

Australia and Tasmania. Vol. 4. (Australian 

Museum: Sydney). 
OAKES, E.J. 1 992. Lekking and the evolution of sexual 

dimoq:hism in birds: comparative approaches. 

The American Naturalist 140: 665-684. 
PRUETT-JONES, M.A. & PRUETT-JONES, S.G. 

1982. Spacing and distribution in Macgregor's 

Bowerbi rd {Amhlyornis macgregoriae). 

Behavioral Ecology and Sociobiology 1 1 : 25-32. 
1983. The Bowerbird's labour of love. Natural 

History 9: 9-54. 
SCHODDE, R. 1976. Evolution in the birds-of-paradise 

and bowerbirds, a resythesis. Proceedings of the 

16th International Ornithological Congress, Pp. 

137-149. (Australian Academy of Science: 

Canberra). 

SCHODDE, R. & MCKEAN, J.L. 1973. Distribution, 
taxonomy and evolution of the Gardener 
Bowerbirds Amhlyornis spp in eastern New 
Guinea witli descriptions of two subspecies. Emu 
73:51-60. 



314 



MEMOIRS OF THE QUEENSLAND MUSEUM 



SCHODDE, R. & TIDEMAN, S.C. (eds) 1988. 

Reader's Digest complete book of Australian 

birds. (Reader's Digest Services: Sydney). 
SIBLEY, C.G & MONROE, B.L. JR. 1990. Distribution 

and taxonomy of birds of the world. (Yale 

University Press: New Haven). 
SIMSON, C.C. 1907. On the habits of the birds-of- 

paradise and bowerbirds of British New Guinea. 

Ibis 1:380-387. 
SOKAL, R.R. & ROHLF, F.J. 1969. Biometry. 

(Freemam: San Francisco). 
STRESEMANN, E. 1953. Laube und Balz der Lauben- 

vogel (Ptilonorhynchidae). Die Vogelwarte 16: 

148-15. 



TRACEY, J.G 1982. The vegetation of the humid 
tropical region of North Queensland. (CSIRO: 
Melbourne). 

VELLENGA, R.E. 1970. Behaviour of the male Satin 
Bowerbird at the bower. The Australian Bird 
Bander 8: 3-11. 

1 980. Distribution of bowers of the Satin Bowerbird 
at Leura, NSW, with notes on parental care, 
development and independence of young. Emu 
80: 97-102. 

WARHAM, J. 1962. Field notes on Australian bower- 
birds and cat-birds. Emu 61: 1-30. 



TABLE 4. Continuity of 51 bower structures at 25 traditional bower sites of male Golden Bowerbirds on the 
Paluma. Range, north Queensland and structural changes over 20 consecutive seasons, from 1 978-1 997. * = AS 
= arboreal subsidiary; TS terrestrial subsidiary; A = arboreal; T = terrestrial; NM = not measured; NC = no 
change; STB = single tower bower; TTB = twin tower bower; RB = rudimentary bower; ** = bowers found 
when under early construction. 



Bower site 
and number 


Number of seasons (=S) 
bower site attended 


When 
examined 


Main tower 


Second tower 


Status 
history * 


Figure 
number 


Height (cm) 


A or T * 


Height (cm) 


AorT* 


la** 


10(S78-S87) 


Mar 79 


25 


A 






STB 


9A 






Apr 80 


NM 


A 






STB 








June 80 


75 


T 






STB 


9B 






Sept 80 


NM 


T 






STB 








Sept 82 


NM 


T 




T 


TTB 








Aug 84 


110 


T 


48 


T 


TTB 




lb 


10(S88-S97) 


Oct 88 


NM 


T 






STB 








Feb 90 


141 


T 


13 


T 


TTB 




2a 


20 (S78-S97) 


Apr 79 


205 


T 






STB 


4E 






Aug 84 


200 


T 






STB 








Feb 90 


198 


T 


10 


A 


STB 








Oct 97 


NM 


T 


15 


A 


TTB 




3a 


14(S78-S81,S84-S93) 


Apr 79 


148 


T 


47 


T 


TTB 


llA 






Aug 84 


NC 


T 


NC 


T 


TTB 








Feb 90 


170 


T 


57 


T 


TTB 


IIB 


3b** 


2 (S82-S83) 


Apr 79 


65 


T 






TS 








May 82 


133 


T 




A 


STB 


9D 






Sept 83 


NM 


T 




A 


TTB 




3c 


4 (S94-S97) 


Oct 95 


NM 


T 




T 


TTB 




4a 


20 (S78-S97) 


Apr 79 


125 


T 


35 


T 


TTB 


5B 






Aug 84 


NC 


T 


NC 


T 


TTB 








Feb 90 


125 


T 


15 


T 


TTB 








Oct 97 


NM 


T 


45 


T 


TTB 




5a 


20 (S78-S97) 


Apr 79 


184 


T 


30 


T 


TTB 








Aug 84 


NC 


T 


NC 


T 


TTB 








Feb 90 


140 


T 


20 


T 


TTB 





BOWERS OF THE GOLDEN BOWERBIRD 



315 



TABLE 4. cont. 



Bower site 
and number 


Number of seasons (=S) 
bower site attended 


When 
examined 


Main tower 


Second tower 


Status 
history * 


Figure 
number 


Height (cm) 




Height (cm) 


A nr T * 

A or 1 


6a 


2 (S78-S79) 


Mar-79 




T 
1 


ID 


T 
1 


TTB 




6b ** 


19(S80-S97) 


Mar-80 


in 


A 






AS 








Oct 80 


JNIVI 


T 
1 


INJVI 




STB 








Sept 81 


MX/I 


T 
1 




A 
r\ 


TTB 








Aug 84 


1 1 n 


T 
1 




T 
1 


TTB 


9C 






Feb 90 


1 Hj 


T 
1 


o / 


T 
i 


TTB 




7a 


18orl9(S78-S95 or 
S96) 


Apr 79 


104 


T 


81 


T 


TTB 








Aug 84 


120 


T 


100 


T 


TTB 








Feb 90 


111 


T 


111 


T 


TTB 




7b 


I or 2 (S96 or S97) 


Oct 97 


NM 


T 


NM 




STB 




8a 


8 (S78-S85) 


Apr 79 


118 


T 






STB 


10c 






Aug 84 


NC 


T 






STB 




8b 


3 (S86-S88) 


Nov 86 


NM 


T 






STB 




8a 


9 (S89-S97) 


Aug 89 


NM 


T 




A 


TTB 








Feb 90 


1070 


T 


55 


T 


TTB 


lOD 


10a 


20 (S78-S97) 


Apr 79 


122 


T 


83 


T 


TTB 


5E 






Oct 85 


NC 


T 


NC 


T 


TTB 








Feb 90 


114 


T 


88 


T 


TTB 




15a ** 


15 (S78-S92) 


Apr 79 


109 


A 






RB 








Sept 84 


NM 


T 






RB 








Sept 88 


NM 


T 


NM 




STB 








Feb 90 


169 


T 


610 


T 


TTB 




15b 


5(S93-S97) 


Dec 93 


30 


T 






STB 








Oct 97 


NM 


T 


20 


T 


TTB 




16a 


8 (S78-S85) 


Apr 79 


150 


T 






STB 


5F 






Aug 84 


NC 


T 






STB 




16b 


3 (S86-S88) 


Nov 86 


BM 


T 






STB 




17a 


8 (S78-S85) 


Apr 79 


107 


T 


98 


T 


TTB 


5D 






Aug 74 


NC 


T 


NC 


T 


TTB 




17b** 


12{S86-S97) 


Nov 86 


NM 


A 






STB 








Oct 87 


NM 


T 






STB 








Feb 90 


168 


T 


121 


A 


TTB 








Oct 97 


NM 


T 


NM 


T 


TTB 




19a 


14{S78-S91) 


Apr 79 


128 


A 






STB 


4D 






Oct 82 


NM 


A 


NM 


A 


TTB 








Aug 84 


110 


A 


30 


A 


TTB 








Nov 86 


NM 


T 


NM 


T 


TTB 








Feb 90 


120 


T 


51 


T 


TTB 




19b 


4 (S92-S97) 


Nov 82 


60 


T 


35 


T 


TTB 








Oct 97 


100 


T 


100 


T 


TTB 




20a 


2 (S78-S79) 


Feb 79 


151 


T 


120 


T 


TTB 


5C 


20b** 


6 (S80-S85) 


Apr 79 


66 


T 






TS 


8A 






Jan 81 


66 


T 


66 




STB 








Sept 82 


NM 


T 


NM 


T 


TTB 








Aug 84 


110 


T 


110 


T 


TTB 




20c 


1 (S87) 


Oct 86 


NM 


T 






RB 





316 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 4. cont. 



Bower site 
and number 


Number of seasons (~S) 
bower site attended 


When 
examined 


Main tower 


Second tower 


history * 


Fi ure 
number 


■1 ICl^iJL V ^111 } 




Height {cm J 


A or T * 


20d 


9 (S89-S97) 


Jan 90 


980 


T 






RB 








Oct 97 


125 


T 






STB 




21a 


5 (S78-S82) 


Apr 79 


124 


T 






RB 


7B 


22a 


1 (S78) 


Apr 79 


143 


T 






STB 


4C 


22b 


13 (S79-S91) 


Sept 79 


1 10 


T 


95 


T 


TTB 


lie 






Aug 84 


170 


T 


130 


T 


TTB 








Feb 90 


174 


T 


95 


T 


TTB 


IID 


22c 


6 (S92-S97) 


Nov 92 


NM 


T 


NM 


T 


TTB 




23 a 


9 {S78-S86) 


Apr 79 


128 


T 






STB 


4B 






Aug 84 


NC 


T 






STB 




23b 


11 (S87-S97) 


Oct 87 


NM 


T 






STB 








Oct 97 


NM 


T 


15 


A 


TTB 




24a 


5 (S78-S82) 


Mar 79 


112 


T 


1 1 1 


T 


TTB 




24b 


6 (S83-S88) 


Aug 84 


125 


T 






STB 




24c 


9 (S89-S97) 


Feb 90 


106 


T 


69 


T 


TTB 




26a 


20 (S78-S97) 


Sept 79 


95 


T 


45 


T 


TTB 








Aug 84 


125 


T 


80 


T 


TTB 








Feb 90 


102 


T 


61 


T 


TTB 




27a 


14(S78-S91) 


Apr 79 


173 


T 


136 


T 


TTB 


HE 






Aug 84 


120 


T 


145 


T 


TTB 








Feb 90 


70 


T 


156 


T 


TTB 


IIF 


27b 


4 or 5 (S92-S95 or S96) 


Nov 92 


60 


T 






RB 




29a 


5 (S78-S82) 


Aug 79 


120 


T 


30 


T 
1 


TTB 




29b 


15 (S83-S97) 


Aug 84 


140 


T 


35 


A 


TTB 


5A 






Feb 90 


140 


T 


1 m 


A 
t\ 


TTB 




33a 


20 (S78-S97) 


Apr 79 


130 


T 


1 Zo 


A 

n. 


TTB 


lOE 






Aug 84 


160 


T 


1 AC\ 


T 


TTB 


lOF 






Feb 90 


141 


T 


1 J J 


T 
1 


TTB 




34a 


16(S78-S93) 


Apr 79 


1 nn 
I uu 


T 
i 






STB 


lOA 






Aug 84 


180 


T 




T 
1 


TTB 


lOB 






Feb 90 


190 


T 




T 
1 


TTB 




34b 


4 (S94-S97) 


Oct 95 


100 


^ 


45 


A 


TTB 


6B 


45a 


1(S78) 


Dec 79 


NM 


T 




T 


TTB 




45b** 


10(S79-S88) 


Dec 79 


50 


A 






AS 








Nov 80 


NM 


A 






STB 








Oct 81 


NM 


T 






STB 








Aug 84 


120 


T 






STB 




45c** 


7 or 8 (S89-S95 or S96) 


Oct 89 


125 


A 






STB 








Sept 90 


160 


T 


80 


A 


TTB 








Nov 92 


NM 


T 


NM 


T 


TTB 




45d 


1 or 2 (S96 or SS97) 


Oct 97 


NM 


T 


NM 


T 


TTB 




47a 


6 {S78-S83) 


June 80 


100 


T 


95 


T 


TTB 




47b** 


2 (S84-S85) 


June 80 


38 


A 






AS 








Nov 84 


165 


T 






STB 




47c 


12{S86-S97) 


Nov 86 


NM 


T 






STB 





CLIFFORD B. FRITH AND DAWN FRfTH 

Frilh* C.B. & Frith. D.W. 2()(..mj Ot> 30; /uicnJancc W\ ch and bciui\-ioLir ai l^ow ci's h\ mak"- 
Gqlden Rowerbifds, Frionodut a noyiomwa (Ptiloflorliv/icludae). Memoirs of (he 
Queensland Mtis^m 4S(2):JJ 7-341. Brisbane. tSISM D{)?9-iH3S. 

Atl«n<idti$& i^el^ ^bwer ilrmalaianpc, bch^vipur of male Golden Bowefbirds 
PribHQ^{tn^pQmanQ at iiw^tte»>vot-siW»'Wbfe Sli«lkle4 jafYerlhe display seasOrtSf of I 
an4 W83 iivStWia of upland rainforest. The clt9pfe(y reason rypically started iti tote- 
August/early Septetnbor and terminated in DcccmbcnJanuarv when wet seiison raiins 
Wmmenced^biitUs length varied ye;»r rci year apparently in response to climate and/or food 
availability. lOuring season 1 9^2 males spenl an average of 36% of daylight at their bowers; 
at .1 mean of 2.7 visits per hour, and each visit averaging 8mins. During season 1 98.^, males 
spent longer iit bowers (mean ^ 6"^%); 'Jl ;i mean of >.4 vi.siis per hour, liiil! each visil 
a\eratiing Mrnins. [,ouer attendanec in I*>S2 in\iW\ecl all males during each muntii and 
dilTerent limes uflhe da>. and \\a> utlnhuted tt' e.\ees;sive!y dr>' eondiii'-ms. Kn\cf aetiv il\ 
ceased by early December, Males spent on average 6% ofihcir lime at bower sites giviiii: 
adverttsemeoi song, 8% other calls. 4^'„ niaimainirig hvtwers. 2% displacement chasing and 
dispia\ ing, and tiie remainnig S0"o perched ^!le^rl^ above their bower. Vocalisations were 
given from hcibiiuall>-u^ed. niosrlv horizontal tM''"i-K perches averaging 'V. froni the 
bower perch and 5.6ni above ground. Ad\erLiscnient sone eon.^isred oK a prolonged, 
pulsating runic repeated an average ofnine times, with each series averaging 33secs- 01 her 
calls included squeak, svr§$ch&:s* wolf-H'/iisllcs, aculds, IVog-audoicada-Jike Ooies, given as 
single notes or as a medley V/&ti fmc vocal avian mimicry of;atl)?ast22 model species. Most 
(95%) boM'cr decorations were collected aw ay from the bowers'iteibavihgbe^ har\ esled, or 
stolen from a neighbouring rival s bower. Olhens w^re r^trieVftd' ftOfll-a- *itore" near the 
bower, wh^t^ they had been left previously. Intruding rival baweeu0WiietsxTWa£ed vo ';teal a 
decoratioB-oniSH of their visits, obvloush^ being jnpst suecessful U00%) tn the owner's 
*b5eitfce.Ofl44displac^mBnt<b^'«»^^ were directed at 

compeciflcs and »H at Mher birds fat lefist ^even spp.). Males instantly displaced and 
ch^iied ail'conspeeirie visitorii f7*^%), mostly from the bower perch. Males were seen to 
display to 3 female-plumaged individual 26 limes. 20 times being beti^re/'atter displacement 
chases. Ttiree display elements w^ere perfonncd by males al bower sites: Bc^w, fleud nod and 
Shake, and i liv^h! Iiovcr. A total of 1 4b displays consisted of one (n ^ 78 1. or a combination of 
(n - 68), these elements, and an elcnienr was often i-epeaied more than once during a display; 
there being no apparent sequence of elements. Copulation wa^ not witnessed. □ Galdetj 
Bowcrhini. Prionodi/n/ newtomona, PtilftpOrhytichUtlBi bOW^T afWsdCdiCe, fime budgets, 
hchcfviow. vtKciiisurinns. 

Clifford B. Frith and Pawn W. Fnih, Honorary Research fd/ows of the Qucvnshnd 
AfiiSewm. ']Pri0nod^a\ B^3SI, Aa/a«5fiT*885. Amrt^a: 17 September J999L 



Until recently fhe^kxltisjaBrtWalJtrdfWorawA^/ti 

Hcwloniana remained One of ftt least studied of 
bowcrbirds. The lirst quanlhafive studies of its 
nesting biology,^ dispersion and constancy of 
bower sitBS, yafifttion and .^eastmal clianges in 
tidw&r stnicture^s, TliDirtie ranges and a8S(}^tated 
Sociobiolo^v and ccologv ha^c only recently 
appeared (Frith & Fnih/|998, 20()Ua. :o00b). 
Male attendance levels at bowers have bect i c ■ - 
anuned io other bowecbird^>6ci6.^ iVeselQv&ky. 
1978; Donaghey. 1981; ?rue<t-J<in^« &. 
PnieH-Jones- 1982 1985; Fnib Frith. 1094; 
Lcnz, 1 993 J, bui until litis study no such data 
vccra avctilable for Gold^ Sowerbfi^ Previous 



contributions prcMtfelcfiialifMlv&taSttai Oba»&r«^ 

aiions of T»iale bower attendance and behaviour, 
but no t|uaniuaii\ c data ( Boiirke & Austin, 1947; 
Marshall. 1954; Chisholm & Chaftcr, 1956; 
Warham, 1962; C balTer. Nis4). 

The display season and male aiiandajice of the 
(jolJcn Bowerbird starts in late August/early 
Sepieniber wtlh bowcr-ovvncrs giving loud, 
pn donged. rattle'ljjte advertiseineni song above 
traditional bowers, adding fresh sticks, acid 
placing decorations upon theiTi (Friih, 1 9S9; Frifti 
& Frith, 2U00a.b|. Other bower calls include 
squeals, screeehe.s, croaks, rasps and churriiigs 
(Schoddc: & Tideiwin, Ftfth. 1989; 



318 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Donaghey, 1996). Males also produce fine vocal 
avian mimicry, as do other bowerbird species 
(Loffredo & Borgia, 1986; Frith & Frith, 
1990a,b; 1993; 1994; Frith & McGuire, 1996; 
Frith et al., 1996). All individually known 
bowerbirds regularly attending and vocalising at 
bowers have proved to be male (Marshall, 1954; 
Vellenaa, 1980; Gilliard, 1969; Cooper & 
Forshaw, 1977; Frith & Frith, 1993). 

Male Golden Bowerbirds spend most time at 
their bower site perched silently above and around 
their bowers, the remaining time being spent in 
calling, displaying, and maintaining and/or 
decorating the bower (Frith, 1989). The bower 
consists typically of one or two roughly conical 
towers of accumulated sticks constructed around 
one or several supporting saplings and/or small 
trees, a perch protruding from single towers or 
connecting twin tower bowers (Frith & Frith, 
2000a). Where tower sticks meet the bower perch 
they are more skilfully placed and ahgned to fomi 
a discrete 'platform ' where bower decorations are 
excusively placed. Frith & Frith (2000a) 
considered the piatform(s) the most significant 
part of the bower structure. For ftirther intro- 
duction, and details ol' structures and dispersion, 
see Frith & Frith (2000a). 

Males display on their bower perch by bowing 
and nodding, with drooped wings, sometimes 
with a bower decoration held in the bill. They also 
display by flying and hovering around the 
immediate bower area (bower site), thus 
dramatically emphasising their brilliant yellow 
plumaae (Chisholm & Chaffer, 1956; Chaffer, 
1958. 1984; Schodde & Tidemann, 1988). 
Copulation has not been observ ed, and may occur 
on or close to the bovver. Males leave their bow er 
site to forage, bathe, collect new bower sticks and 
harvest, or steal, decorations. That male bower- 
birds steal decorations trom the bowers of rivals, 
with a preference for particular colours and items, 
has long been known (Marshall, 1954 and 
references therein), but has been only briefly 
alluded to with respect to Golden Bowerbirds 
(Frith, 1989). It has been described for several 
bowerbird species (Borgia, 1985b,c, 1986; 
Borgia & Gore, 1986; Pniett-.lones & Pruett- 
Jones, 1994; Frith & Frith 1993, 1994, 1995; 
Hunter & Dwyer, 1997). 

In this contribution, we initially define and 
describe seasonality of bower attendance over the 
first three display seasons of our study ( 1 978-80) 
in relation to rainfall, temperature and fruit and 
insect food availability. However, most data 



presented here deal with bower site attendance 
levels of males over two display seasons 
(1982-83), diurnal, monthly and seasonal vari- 
ations in these, and behaviour and vocalisations 
at the bower. We discuss these results in the 
context of knowledge of this and other 
bowerbirds. 

METHODS 

STUDY AREA AND CLIMATE. The main 
study area comprised 50ha of upland tropical 
rainforest, at about 875m asl, on the Paluma 
Range (19°00'S, 146°10'E), northeastern 
Queensland, 7km from Paluma Township and 
80km north of Townsville. This area, measuring 
1 X 0.5km, was permanenllv tiridded w ith metal 
stakes (see Frith & Frith, 2000a: fig. 2). The 
rainforest has been classified as simple nolophyll 
vine forest (Tracey, 1982). 

Annual rainfall and temperature show marked 
seasonality on the Paluma Ran^e (Frilh, 1984; 
Frith & Frith, 1985, 1994; D. Fnth & C. Frith, 
1990). The dry season extends from April- 
November, with June-August the driest and 
coldest months. Rainfall and temperatures 
increase during September-October and decrease 
during April-May. The hotter wet season is from 
December-March, with most rain falling during 
January-March. 

DEFFNITIONS. Bower site describes the location 
of a traditional bower; regularly attended, 
maintained and decorated throughout each 
season by the traditional adult male owner (Frith 
& Frith, 2000a). A traditional bower owner was 
an individually-marked (colour-banded), 
bower-attending, bird known to have attend a 
particular bower during at least one previous 
season(s). Male attendance refers to known 
individual males perching, calling, displaying at 
or maintaining their own bower. Thus a male 
visiting the bower of another to steal a decoration 
was not attending it. As we could see only the 
male bower-owner during most displays we refer 
to them as 'displays', as distinct from 'courtship 
displays' (i.e. display directed at a conspecific). 
We use 'regularly attended' to imply frill-time 
seasonal attendance by traditional owners at 
traditional bower sites, and write about males 
unless stated otherwise. To *har\^est' a bower 
decoration was to obtain it from a plant or the 
forest floor, as distinct from stealing it from the 
bower ofarival male. We refer to a display season 
by the year in which it started (S78, S79 etc). 



BOWER ATTENDANCE AND jBEHAVlOURBY GOLDEN BOWERBIRDS 319 




flQ. I. Monthly rainfa]! (= colunins) from August TS^8-December I985vJi*CQrpQrating.fTve displity seasons 
(S78-S53)^ mi monW^^^m^f^ 191B^] 990 (= ■) on the P^lumft Iteoge, noith Queenstiria. 



ANNUAL SEASONALITY OF BOWER 
ATTENDANCE BY MALES. Seasonality of 
bower attendance was assessed during S78-SS1 
by (a) ^stunating numbers of bovv er decorations 
on bowers at our visits (see Frith & Frith, 
ZOOOa.b) and subjeciivcly calcgDrising ihem as 
being poorlv (<10 decorations), moderatelv 
(t0-2ffy0r Well (>20) decorated; (b)Ti0tiiig how 
JiELa^y finaieE^ W ]>£ard advettisaKL^ SQt^s at 
te^dttitjtisd bdiwer sites diiring 28Cih tyf trani^ 
foraging walks from August V'H^ to Februarx 
1981; and (c) collecting defaecated seeds on 
black niesb cjtchment traps suspended beneath 
favoured perches aboVp or adjacent to, up to ten, 
bowers at regular 'fiitervals during December 

1978- May 1979 and Sq;rteii?ber 1 979-Febniaiy 
1981. 

jScrasonaiity bow^ antendance during 

S78-S8i Was examined in relation to rainfall 
(Fig. 1). temperature, relative irint crop and 
itisect numbers (Fig. 2). We collected tree fruiting 
pheqology data iroru 602 trees during September 
197«-A33Til 1979, ^thereafter about 500 of 
-the^e tteeji were exaltiitted at six- (July 

1979- Atigust 1980) or eight- (November 
19X0-Fcbruarv i^Sl) weekly uitervals (Frith 
Fruh I9Q4). Diurnal msect populations 
were monitored each month from August 
1978-April 1979, and July 1979-Fsbmary 19S1,. 
usitig Malaise traps (Frith & Frith, 19S5). Wfr 
present here the mean diurnal number of all 
insects trapped per month, and for Coleoplera 
separatdy because 80% of any animals xemaiiis 



f o u n d 



in 



'a c c a 1 samples during A u gust 



1978-F6bniaQ' 1981 were coleopteran (Fridi & 
Fritfi, unptiH data). 

M.VLL LV.)\VFR ATTLNDANCF LFVELS. 
Male ho^ser attendance levels were monitored 
during the peak display period of 7 September- 1 5 
November 1982. Season 82 was exceptionally 
dry aiid bower attendance d^€9^cl iPQQSi^f^r-* 
ably by early Novetnbef. We therefofe repeat^ 
obser\'ations the following season, during ? 
November-5 December, when rainfall was only 
just below a\ erage, and bowers were regularly 
attended. Observations ovet iwo seaports 

and diumaT<vat^cVi5. 

adjacent bowerSc ni^re nldnitorgd during 
bcrfh seasOfK fbpwifrs 1, 2, 5, 4, ^4iavi ilh 5?e 

Frith & Frith, 200na: Jig, 2). We esta^&l|ed' 
eiyptic canvas hides six metres from eaeh"boWei* 
tv^'o weeks before starring observations. Each 
observation lasted six unmterrupied hours, 
during 0600-1200 or 1200-I800h, over peak 
seasonal activity. Each cycle of observatioiis 
consisted of two (at 0600-1 200b and at 
1 200-1 SOOhl periods at each of the six bowers 
(dius 12 X oil obsen ations). When a cycle was 
cornpleted w e repeated it. Observation periods at 
tfatPR^efe of two adjacent bowers (bowers 1 an^ 3 
at 2gl)ni apart; 2 and 4 at 2 1 Om apart; 1 9 artdSO at 
130m apart) were made simultaneously by us, 
D WF in one hide and CBF in another. During SS2 
we made 1 50, L47'Bnd 72fe of direct observalion 



320 



MEMOIRS OF THE QUEENSLAND MUSEUM 






FIG. 2. A, mean monthly temperatures (= m) and percentages of trees (= columns) sampled monthly that were in 
fruit. B, mean monthly numbers of all diurnal insects (= columns) and of only Coleoptera (= ■) sampled by 
Malaise traps. C, mean monthly numbers of defaecated seeds (= columns) collected beneath Golden Bowerbird 
singing perches and the number of advertisement songs (= ■) heard during track transect walks (see Methods), 
from August 1978-February 1981 and incorporating three display seasons (S78-S80), on the Paluma Range, N 
Queensland. 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



TABLE L Bower site attendance levels by individual male Golden Bowerbirds during the display seasons of 
1982 (September, October and November) and 1983 (November). * = minimum number for males at bowers 2, 
4, 19 and 20 who may have been at that site pre-S78; ** = immature plumage in season 82, adult plumage in 
season 83; *** = a different adult male regularly attended this bower site during the first two September 
watches; **** = a different adult male each season; see Methods. 





Bower site 
number 


Number of 


Hours and 


Absence 


Presence 


Season 


seasons owned 
* 


(numbers) of 
obsen'ations 


Mean mins per 
absence 


% of total time 
absent 


Mean mins per 
presence 


% of total time 
present 


1982 


] 


5 




12.1 


67.9 


5.9 


32.1 




2 




60 { 1 0) 


13.8 


67.1 


6.9 


32 9 




3** 


1 


60(10) 


16.2 


66.4 


8.6 


33.6 




4*** 


5 


60(10) 


11.6 


53.0 


10.4 


47.0 






5 


66(12) 


13.0 


63.5 


7.6 


36.5 




20 


5 


63(11) 


17.3 


66.9 


9.1 


33.1 


TotaI/Mean/% 






357 (61) 


13.9 


64.0 


8.0 


36.0 


1983 


1 


6 


12(2) 


7.8 


47.5 


8.2 


52.5 




2 


6 


24(4) 


5.5 


38.8 


8.8 


61.2 




3** 


2 


12(2) 


7.8 


32.6 


16.7 


67.4 




4 


6 


18(3) 


5.2 


35.0 


9.6 


65.0 




19*** 


1 


18(4) 


9.6 


38.8 


14.2 


61.2 




20 


6 


18(4) 


6.3 


31.4 


13.5 


68.6 


Total/Mean/% 






102(19) 


6.6 


37.0 


11.0 


63.0 



during September, October and November, 
respectively. Fieldwork terminated in November 
due to extremely dry conditions resulting in males 
irregularly attending bowers. During S83 we 
made 90h of direct observation during Nov- 
ember, until heavy rains hampered fieldwork. 
The last 12h observation cycle in November had 
to be postponed until 5 December, but December 
data are combined with November results herein. 
Fieldwork then ceased because continuing 
torrential rains resulted in males irregularly 
attending bowers. 

To analyse diurnal variation we subdivided the 
totals for male attendance levels into four periods 
(0600-0900, 0900-1200, 1200-1500, 1500-I800h). 
This made data directly comparable with a 
similar study of Tooth-billed Bowerbirds 
Scenopoeetes dentirostris (see Frith & Frith, 
1994). We used the same periods to analyse 
vocalisation frequencies (see below). 

Determining actual time a bower-owning male 
spent at his bower site was often difficult. Most 
limes we saw an absent male return by flying to 
one of his favoured perches, or onto his bower 
perch. However, sometimes the first indication of 
his renewed presence was when he called. If he 
remained out of sight but continued to call we 
assumed he was present, especially if later he 
flew to another perch, to the bower, or away from 
his bower site. If we did not re-sight or hear him 



we considered his time present to be ended at his 
last recorded call, even though he may have 
subsequently remained above the bower for some 
(limited, in our experience) time. We usually saw 
the male fly off, but sometimes he would fly 
unseen higher into the canopy and we were 
unsure if he had left, unless he gave progressively 
distant vocalisations as departing. Thus, times 
presented in Tables 1-3 for males spent at their 
bower sites are minimums. Having said that, the 
times we recorded each of the six males at their 
bower sites were similar each season. This 
suggests that any discrepancy between the time 
we recorded present and the actual time involved 
may be minimal. Single call notes (see below), 
occasionally heard some distance (>30-40m) 
from bower sites during a male's apparent 
absence, were discounted as indicative of his 
presence, as we could not confirm they were in 
fact given by the bower owner. 

BOWER OWNERSHIP. Males were mist-netted 
at or near bowers and marked with a metal 
Australian Bird and Bat Banding Scheme band 
and a unique two colour band combination (= 
marked), and released at the capture location. 
Banded males included not only the owners of the 
six bowers under intensive observation, but also 
males intruding from adjacent bowers. 

Males attending four of the six bowers in S82 
had regularly attended their respective bowers as 



322 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. Monthly bower attendance by male Golden Bowerbirds during September, October and November of 
the display season of 1982. 



Months 


Bower site 
number 


Hours and 
(numbers) of 
observations 


Absence 


Presence 


N'tean mins per 
absence 


% of total time 
absent 


M^ean mins per 
presence 


% of total time 
present 


September 
Total/Mean/% 


1 


12(2) 


13.6 


75.3 


4.6 


24.7 


2 


24 (4) 


18.6 


72.5 


7.2 


27.5 


3 


24 (4) 


18.3 


67.2 


9.6 


32.8 


4 


24 (4) 


15.6 


48.8 


16.4 


51.2 


19 


27 (5) 


14.7 


60.0 


9.5 


40.0 


20 


27 (5) 


16.9 


57.2 


13.6 


42.8 




138(24) 


16.4 


62.0 


10.3 


38.0 


October 
Total/Mean/% 


I 


24 (4) 


11.2 


62.0 


7.0 


38.0 


2 


24 (4) 


11.4 


62.6 


6.8 


37.4 


3 


24(4) 


!3.4 


61.5 


8.5 


38.5 


4 


24 (4) 


10.0 


54.6 


8.5 


45.4 


19 


27 (5) 


10.4 


63.1 


6.4 


36.9 


20 


24 (4) 


16.1 


70.2 


7.0 


29.8 




147 (25) 


11.8 


62.0 


7.3 


37.6 


November 
Totaf/Mean/% 


1 


12(2) 


12.7 


72.2 


5.0 


27.8 


2 


12(2) 


11.8 


65.3 


6.4 


34.7 


3 


12(2) 


18.5 


74.4 


6.8 


25.6 


4 


12(2) 


10.5 


58.1 


7.4 


41.9 


19 


12(2) 


17.9 


72.1 


7.2 


27.9 


20 


12(2) 


21.1 


82.2 


4.9 


17.8 




72(12) 


14.8 


71.0 


6.3 


29.0 



meaningfully, and so we estimated their totals by 
allowing 2secs for each. Calls given during bower 
maintenance, displacement chases and display 
periods were, however, included in time periods 
totalled for those activities. The number, 
behaviour and vocalisations of female-plumaged 
and adult male visitors/intruders to bowers were 
monitored and their presences timed. Numbers of 
decoration thefts by rival bower-owners were 
recorded. Numbers, distances from bowers, and 
heights of habitually-used perches were noted. 
The length of time favoured perches were used at 
bower sites 2, 4 and 1 9 during S82 was recorded. 

Bower maintenance periods included time a 
male was on the bower perch and adding a 
decoration or a stick to it, or adjusting and/or 
tidying. Most visits to the bower perch were 
solely for maintenance, but some were exclus- 
ively to display. Sometimes a male displayed on 
the bower perch before starting maintenance. In 
the latter case each behaviour was timed 
separately. Similarly, when a display was 
instantly followed by a displacement chase both 
periods of behaviour were treated separately 
even when directed at the same visitor/intruder. 



aduh-plumaged individuals since at least S78. In 
S83 three of them (at bowers 2, 4 and 20) 
remained in attendance, but the male at bower 19 
had been replaced by another male we tlrst 
caught (at bower 19) in adult plumage in May 
1982. The male attending bower 1 in S82 was 
first caught as an immature in March 1 979, when 
he had Just taken the site over and was building a 
new bower there. He acquired adult-plumaged in 
S80. The male attending bower 3 was still 
immature (female-plumaged) during the first 
season (S82) of this study. We first banded him in 
March 1979, at a point 140m from bower site 3. 
He attained adult-plumage during the second 
season (S83) of this study. 

MALE BEHAVIOUR AT BOWERS. Male 
behaviour at bowers was categorised as: periods 
of advertisement song, or other calls (including 
single notes and medleys); bower maintenance; 
displacement chases; displays; and silence. 
Advertisement song and other calls were given 
from favoured perches above or within 15m of 
the bower, and were timed and totalled separately 
as they involved no other behaviour. Single calls 
were too brief (mostly <2secs) to time 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



323 



TABLE 3. Variation in diurnal bower attendance levels of male Golden Bowerbirds during the display seasons of 
1982 and 1983. 



Season/time 


Month 


Hours and 
(number) of 
obser\'ations 


Absence 


Presence 


Mean mins per 
absence 


% of total tmie 
absent 


Mean mins per 
presence 


/o or loiai tmie 
present 


1982 

0600-0900 
Total/Mean/% 


September 


33(11) 


9.8 


57.3 


7.4 


42.7 


October 


36(12) 


7.4 


51.6 


7.0 


48.4 


November 


18(6) 


9.0 


62.3 


5.6 


37.7 




87 (29) 


8.6 


56.0 


6.8 


44.0 


0900-1200 
Total/Mean/% 


September 


39(13) 


11.5 


49.5 


11.7 


50.5 


October 


39(13) 


12.3 


59.3 


8.7 


40.7 


November 


18(6) 


13.8 


65.0 


7.9 


35.0 




96 (32) 






O 7 


4j.O 


1200-1500 
Total/Meaii/% 


September 


33 (11) 


24.9 


69.2 


1 T A 
I —.4 


in B 


October 


36(12) 


15.5 


71-8 


0.4 


TOT 


November 


18(6) 


21.7 


82.4 


5.1 


17.6 




87 (29) 


19.4 


73.0 


7.8 


27.0 


1500-1800 
Total/Mean/% 


September 


33 (11) 


29.1 


74.9 


10.6 


25.1 


October 


36 (12) 


12.4 


67.1 


6.4 


32.9 


November 


18(6) 


16.1 


73.1 


6.2 


26.9 




87 (29) 


17.2 


71.3 


7.3 


28.7 


1983 

0600-0900 


November 


30(10) 


5.5 


34.2 


10.4 


65.8 


0900-1200 


November 


30(10) 


6.2 ' 37.2 


10.2 


62.8 


1200-1500 


November 


25 (9) 


8.4 


44.3 


10.6 


55.7 


1500-1800 


November 


17(6) 


6.1 


31,7 


12.5 


68.3 



Each period of display included one to several 
display elements. A display element consisted of 
any one of the three distinct displays perfomied 
by male Golden Bowerbirds. 

Chi-squared tests and Student's two-tailed 
/-tests were used for statistical comparisons. 
Percentage data were normalised by applying 
arcsin transfonnation. Means are given as ± one 
standard deviation. In some instances we also 
present standard error, to facilitate comparisons 
with data presented by other bowerbird studies. 

RESULTS 

ANNUAL SEASONALITY OF BOWER 
ATTENDANCE BY MALES. Regular seasonal 
attendance of traditional sites and bowers, by 
their traditional owners, typically started on the 
Paluma Range in late August/early September. 
The commencement, length, and termination, of 
a display season varied from year to year, 
primarily in response to climate and/or fruit 
phenology, as illustrated by results of the first 
three seasons of our study (August 1978 to Feb- 
ruary 1981). 



We recorded the seasonally first bower ad- 
vertisement songs during 14-17 August at the 
start of S78, and by 2 1 August some bowers had a 
few decorations on them. By early September 
most bowers were moderately decorated, with 
new sticks added to them. From the second week 
of September until the end of December bower 
sites were regularly attended by their traditional 
owners. Rainfall was slightly above average for 
the time of year (Fig. 1 ). Temperatures increased 
during these months, from an average of 1 9°C in 
September to 25°C in December, and fruits and 
insects were plentiful (Fig. 2A, B). Bowers 
remained moderately decorated and attended 
until the end of December, but then activities 
decreased as rainfall increased. During the last 
week of January, 594mm of rain fell and bower 
attendance ceased. Rain continued throughout 
February, to 15 March, as bower decorations 
deteriorated. No advertisement song was heard, 
but limited faeces beneath favoured perches 
indicated some males had briefly visited bower 
sites (Fig. 2C). During brief dry spells, one or two 
fresh decorations were sometimes placed on 
bowers. There was then a brief period of renewed 
activity during late March to the first week of 



324 



MEMOIRS OF THE QUEENSLAND MUSEUM 



May, but bowers were poorly decorated and few 
advertisement songs given (Fig. 2C). 

Bowers were undecorated/unattended during 
June/July 1979, and not until 19 August did we 
hear the first advertisement song, marking the 
commencement of S79. During September 1979, 
rainfall was average for the month and temp- 
eratures rose, but the fruit crop was sparse and 
remained so throughout the display season ( Figs 
1, 2A). Insects, including Coleoptera, were less 
abundant than the previous season (Fig. 2B). 
October and November were exceptionally dry 
and hot with rainfall (64mm) well below the 
seasonal average (230mm), During November, 
fewer advertisement songs were heard and, while 
bowers were poorly/moderately decorated, 
faecal samples indicated males were attending 
bowers if not maintaining them (Fig. 2C). It 
remained dry until 25 December; by which time 
bower attendance had declined, few calls were 
given, and bower decorations dried and were not 
replenished. It rained heavily from the last week 
of December until 12 March, with little or no 
bower attendance. As in the previous year, there 
was renewed activity during March, as rains 
eased, that lasted until about the second week of 
May. 

There was a notable increase in available fruit 
crop during winter months of 1980 (Fig. 2A). 
Some bower owners placed a few decorations on 
traditional bowers by mid-June-July, started 
advertisetnent song, and accumulating faeces in- 
dicated males were now spending time at bowers 
(Fig. 2). This winter attendance continued 
through to August, possibly because of a larger 
fruit crop. By August 1980 all bowers were 
regularly attended, despite lack of rain (Fig. 1). 
Temperatures increased notably in September, 
insects were abundant, and fruit plentiful; and 
bowers were well attended as indicated by faeces 
at them (Fig. 2). Rainfall during September- 
December was near seasonal average, but from 1 
Januar\' was excessive, falling e\ery day until 
26th (2201mm; see Fig. I). Bower decorations 
deteriorated during January and, while no 
advertisement song was heard, faeces indicated 
males occasionally visited bower sites (Fig. 2C). 

We did not monitor bower activities as closely 
over the next three years, but seasonal trends 
showed a similar pattern with regard to relative 
rainfall. In S8 1 bower activities commenced during 
mid-August and lasted until mid-November 
when, due to heavy rain (468mm), they slowed 
and then ceased in December (Fig. 1). During 



October and November of S82 it was exception- 
ally dry (7lmm), and bower activity levels were 
similar to those described for S79 (see above). 
The display season commenced earlier the 
following season, much as for S80. In S83 
rainfall was near the seasonal average, and bower 
activities persisted until the commencement of 
the January rains. Bower attendance levels, and 
behaviours and vocalisations at traditional 
bowers during S82 and S83, are discussed in 
detail below. 

RELATIVE LEVELS OF BOWER ATTEND- 
ANCE BY MALES. Seasonal variation. During 
S82 males spent an average of 36% of total time 
at their bowers (Table 1), proportional differ- 
ences between individuals not being significant 
(X^ = 4.46, P>0.30.). During S83 males spent an 
average of 63% of total time at their bowers 
(Table 1), proportional differences between 
individuals likewise not being significant (x^ = 
2.72, P>0.70). Males spent an average of 8 (SE = 
0.8) and 11 (SE 1.3) mins at bowers per 
presence, and absences averaged 13.9 (SE = 1 .7) 
and 6.6 (SE = 0.7) mins during S82 and S83 
respectively. Mean number of visits per hour was 
2.7 (range 2.2-3.3) and 3.4 (range 2.6-4.2) during 
S82 and S83 respectively. Thus, all males 
attended their bowers for far less (27%) time, less 
frequentlv, and for less time per visit, during S82 
than during S83 (Table I). 

Monthly variation. During S82 there was a 
significant difference between the proportion of 
total time individual males spent at bowers 
during September (x^ = 13.74, P<0.02), because 
not all started attending bowers at the same time 
and/or with the same intensity (Table 2). At 
bower I, the owner was not sighted on 13 
September (the first S82 observation), but two 
immature males were briefly (<5% of 
observation) seen adjusting its decorations and 
sticks. These young males gave occasional 
screech and scold notes near the bower, but no 
adverfisement song. No birds were here on 14 
September but at our next observation, on the 
27th, the traditional owner was regularly 
attending; but at a mean duration per presence 
lower than other males (Table 2). 

At bower 4, male attendance was notably high 
in September 1982 (Table 2); apparently because 
a male new to it, in his first year of adult plumage, 
was regularly attending (51% of total time) on 9 
and 10 September (the first two S82 observ- 
ations). We assumed he was the new owner, but 
during our next two observafions, of 21 and 22 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



325 



FIG. 3. The percentage of total time that each of six aduh male Golden Bowerbirds spent attending their 
traditional bower site, during bi-weekly periods of the display seasons of 1 982 ( 1 Sept.- 1 5 Nov. 1 982) and 1 983 
(Nov. I- Dec. 15), relative to the amount of rain (column) that fell during each period. Symbols indicate the 
bower sites: t = 1, □ - 2, • =3, ■ - 4, o = 19, u = 20 (see Tables I & 2). 



September, the traditional owner was in regularly 
attendance (50% of total time); presumably 
having displaced the challenger. Mean duration 
per presence of the challenger (28.9 mins) during 
the first two September observations was far 
greater than that of the owner ( 1 1 .4 mins) during 
the latter two observations, and was greater than 
that of other individuals during September or any 
other month (Table 2). Male attendance at bower 
4 remained relatively high throughout S82. 

September S82 rainfall was average, but 
October was exceptionally dry and hot (only 
6mm of rain, on the 4th; Fig. 1 ). Male bower 
attendance levels increased little during October 
over those of September, actually decreasing 
slightly in the middle of the month, with mean 
duration per presence lower (Table 2, Fig. 3). 
There was no significant difference between the 
proportion of total time individual males spent at 
bowers (x- = 3.54, P>0.50). November rainfall 
(68mm) was well below the average (157mm), 
the first two weeks being particularly dry 
(2 1 mm). By mid-November bowers were poorly 
maintained and decorated, few advertisement songs 
were given, and attendance levels decreased 
considerably (Table 2, Fig. 3). There was a sig- 
nificant difference between time individual males 
spent at bowers during November {x~ = 11.55, 
P<0.05), because they stopped attending bowers 
at different times (Table 2). Despite much more 



rain in December, attendance levels did not 
recover, males were rarely sighted at bowers. 

In S83 rainfall was near average (Fig 1). 
Although we made observations only during 
November 1983, bower attendance by all in- 
dividuals was much higher (63%) than in 
September (38%), October (38%) or November 
(27%) of S82 (Tables 1, 2; Fig. 3). There was no 
significant difference between the proportion of 
total time individual males spent at bowsers (x2 = 
2.72, P>0.70) in November of S83. 

Diurnal variation. During 0600-0900, 0900- 
1200, 1200-1500 and 1500'"-1800h ofS82 males 
spent 44, 44, 27 and 29% of total time attending 
bowers respectively, differences between these 
proportions being significant (x^ = 7. 1 5, P<0. 1 0). 
Thus males spent much more S82 time attending 
bowers in mornings than afternoons, a trend 
apparent during September, October and 
November (Table 3). 

During the same four diurnal periods of S83 
males spent 66, 63, 56 and 68% of total time 
attending bowers respectively, differences 
between these not being significant (x" = 1.41, 
P>0.70). Thus, male attendance levels at bowers 
were much higher in S83 than in S82, both in 
mornings (by 21%) and afternoons (by 34%). 
Mean duration per bower attendance was higher 
throughout the day in S83 than in S82 (Table 3). 

MALE BEHAVIOUR AT BOWERS. Habitual 
perches. Males had several favoured perches 



326 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 4. Time-budgeted activities performed by individual male Golden Bowerbirds at bower sites during the 
display seasons of 1982 (September, October and November) and 1983 (November). * = immature male during 
season 82; ** ^ data for bower 3 excluded; see Results. 





Advertisement song 


Other calls 


Maintenance 


Smgle 


Medley 


Season 


Bower site 
number 


Mean sees 
per song 


Mean no. 
rattles per 
song 


% of time 
present 


% of time 
present 


Mean sees 
per call 


% of time 


Mean sees 

Ud L>C1 IVJU. 


% of time 

Ui C^CI Jl 


1982 

Total/Meaii/% 


1 


38 


8 


9.1 


I.l 


105 


3.6 


61 


7.8 


2 


39 


7 


7.1 


1.5 


121 


3.9 


70 


7.1 


3 * 


36 


10 


5.1 


2.0 


279 


34.5 


63 


5.8 


4 


32 


9 


5.4 


1.0 


125 


4.3 


48 


4.9 


19 


37 


6 


6.2 


1.0 


119 


2.9 


56 


5.3 


20 


35 


7 


6.5 


1.0 


107 


4.8 


62 


4.7 




36 


8 


6.4 


1.2 


177(111**) 


8.7(5.4**) 


59 


5.8 


1983 

Total/Mean/% 


1 


31 


9 


7.8 


0.4 


72 


2.9 


62 


3.0 


2 


31 


9 


4.4 


0.2 


101 


4.6 


45 


3.5 


3 * 


33 


9 


6.8 


0.4 


110 


6.1 


20 


0.6 


4 


34 


10 


4.9 


0.3 


63 


2.7 


43 


3.6 


19 


28 


8 


4.6 


0.7 


143 


15.4 


37 


5.4 


20 


27 


7 


4.2 


0.4 


106 


3.6 


22 


0.5 




30 


9 


5.1 


0.4 


110 


5.9 


40 


2.8 






Displacement chases 


Display 


Silence 


Total time 
present 
(mins) 






Season 


Bower site 
number 


Mean sees 
per chase 


% of time 
present 


Mean sees 
per display 


% of time 
present 


% of time 
present 






1982 

Total/Mean/% 


1 


34 


0.9 


59 


1.6 


75.9 


925 






2 


11 


0.2 


44 


1.5 


78.7 


1185 






3 * 


20 


0.4 


60 


0.9 


51.3 


1210 






4 


50 


1.0 


66 


1.0 


82.4 


1693 






19 


20 


0.2 


64 


0.8 


83.6 


1444 






20 


34 


1.1 


48 


1.0 


80.8 


1250 








31 


0.6 


58 


1.1 


76.2 


7707 






1983 

Total/Mean/% 


1 


36 


l.I 


76 


1.3 


83.5 


378 






2 


39 


0.4 


35 


1.2 


85.7 


881 






3 * 


43 


1.3 


60 


1.0 


83.8 


485 






4 


47 


1.5 


51 


1.8 


85.2 


702 






19 


31 


0.6 


36 


0.8 


72.5 


665 






20 


21 


0.3 


35 


0.6 


90.4 


741 








38 


0.8 


43 


1.1 


83.9 


3852 







above and around their bower, on which they 
gave advertisement vocalisations, perched 
silently, or preened. Of a total 947 occasions (S82 
and S83 combined) that males were recorded 
perched above/around the bower, 99.6% 
involved horizontal branches, mostly of saplings 
or small trees, and the remainder horizontal to 
gently sloping vines. Where some of these 
horizontal branches abutted the plant's vertical 
trunk (5% of perches used), males sometimes 



placed sticks to form small arboreal subsidiary 
bower structures. During S82 males used an 
average of 11.7 ± 3.9 perches per observation 
period (n = 61), at a mean of 4.9 ± 2.5m above 
ground, and 9.5 ± 2.6m distant from the bower 
perch. During S83 these figures were 13.1 ± 
4.1m, 6.3 ± 1.9m and 9.0 ± 2.6m (n =19) 
respectively. During S 82 males at bowers 2, 4 and 
19 spent 15% of their time perched above their 
bowers on perches known to be favoured ones. 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



327 



Advertisement song. This consisted of a pro- 
longed pulsating rattle note, t>'pically lasting one 
or two seconds, that was usually repeated a num- 
ber of times. It was difficult to precisely locate a 
bird giving this call, and others. Occasionally 
males gave only one rattle, or a series of one to 
three, when first arriving back at the bower and 
before commencing a much longer song (up to 27 
rattles being recorded). Number of rattles per 
song averaged eight in S82 and nine in S83, each 
song averaging 36 and 30 sees respectively 
(Table 4). During some visits males only main- 
tained bowers, and did not give advertisement 
song, whereas during others they gave several 
sets of rattle song (10 songs being the most 
during a single visit). Males gave advertisement 
song on 48% of 1 706 bower visits in S82, and on 
61% of 645 visits in S83. 

Males spent 6.4 and 5A% of time present at 
bowers giving advertisement song during S82 
and S83 respectively (Table 4). There was no 
significant difference between the proportion of 
time individual males spent giving advertisement 
song at their bowers each season (82: X" - 1-57, 
P>0.90; 83: = 2.02, P>0.80), nor were dif- 
ferences between them for the two seasons 
significant (arcsin transfonnation t|Q = 1.39, 
P>0.20). Mean duration of each song period was 
similar diu^ing each month of S82 (Tables 5 & 6). 

During September of S82, as males re- 
established themselves, individuals spent more 
time at bowers giving advertisement song (7.5%)) 
than in October (5.9%) and November (5.0%). 
Mean duration of each song was also longer 
(Table 5). The male at bower I did not start bow er 
attendance until late September, but was par- 
ticularly vociferous (Table 5). The lower S82 
October and November tlgures may have 
reflected extremely dry conditions; but in S83, 
when climate was more favourable, males still 
spent only 5.1% of their presence giving 
advertisement song (Table 4; Figs 1,2). 

Males spent more of their presence at bowers 
giving advertisement song during mornings than 
afternoons during S82 and S83 (Table 6); pro- 
portional differences between diurnal periods (data 
for both seasons combined) being significant 
(arcsin transformation, t4 = 5.33, P<0.01). Mean 
duration of songs was similar at different times of 
day (Table 6). In S82 males performed 37, 43, 1 1 
and 9% of songs (n = 819) during 0600-0900, 
0900-1200, 1200-1500 and 1500-1 800h respect- 
ively. In S83 they performed 46, 27, 14 and 13% 
of songs (n = 391) during the same periods 
respectively. 



Other calls. These consisted of single calls or a 
continuous medley of them. Single ones were a 
squeal, screech, scold-rasp, or wolf whistle as 
follows: squeal was a high-pitched thin and 
variable note; screech a variable, harsher and 
lower, but louder and more assertive, note 
sometimes delivered with a rather braying-like 
quality; scold-rasp a loud and urgent note(s); and 
wolf-whistle a powerful, two note, harsh and dr}', 
squeal/screech notes with the same cadence and 
timing as a human 'wolf-whistle'. 

Single calls, such as screech and wolf-whistle, 
were mostly given as males approached or left 
their bower site, or when conspecifics w ere close. 
They were also heard some distance away from 
bowers, while owners were absent and presum- 
ably foraging. The scold-rasp was sometimes 
given when an inter-specific bird, larger than the 
bower-owner, such as a Spotted Catbird, 
Ailuroedus melanotis, or Satin Bowerbird, Ptilo- 
norhynchus violaceus, came close to or onto the 
bower. Once when an Australian Brush-turkey, 
Alectura lathanh, walked over a bower the male 
owner scolded it for 60secs, until it left. 

During S82 and S83 males gave single calls for 
1.2% and 0.4% of time present at bowers, 
respectively (Table 4). The proportion of single 
calls given was broadly similar each month of 
S82 (Table 5), and for different times of the day 
(Table 6), during both seasons. 

A medley included a continuous series of single 
calls interspersed with frog- and cicada-like notes, 
a single rattle with a squeal(s) and/or vocal avian 
mimicry. At least 22 bird species were identified 
as models for mimiciy perfonned: White-headed 
Pigeon, Columha leucomela\ Red-tailed Black 
Cockatoo, Calyptorhynchus hanksii\ Sulphur- 
crested Cockatoo, Cacatua galerita\ Australian 
King-Parrot, Alisterus scapularis; Crimson 
Rosella, Platycercus elegans; a cuckoo; Noisy 
Pitta, Pitta versicolor; Yellow-throated Scrub W- 
ren, Sericornis citreogularis\ Large-billed 
Scrubwren, S. magnirostris; Brown Gerygone, 
Gerygone mouki; Mountain Thombill, Acanthiza 
pusilla; Bridled Honeyeater, Lichenostomus 
frenatus; Grey-headed Robin, Heteromyias 
albispecularis; Chowchilla, Orthomvc spaldingii; 
Eastern Wliipbird, Psophodes olivaceus; Bower's 
Shrike Thrush, Colluricincla boweri; Barred 
Cuckoo-shrike, Coracina lineata: Pied Currawong, 
Strepera graculina: Victoria's Rifiebird, Ptiloris 
victoriae; Spotted Catbird; Tooth-billed Bower- 
bird; Satin Bowerbird and many small passerine 
notes we did not identify. 



328 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 5. Monthly time-budgeted activities performed by individual male Golden Bowerbirds at bower sites 
during September, October and November of the display season of 1 982. * = immature male during season 82; 
** = % of time present with data for bower 3 excluded; see Results. 







Advertisement song 


Other calls 












Medley 


Mamtenance 


Months 


Bower site 
number 


per song 


/Q KJl LililC 

present 


/o oi lime 
present 


Mean sees 
per call 


/O Ui LllliC 

present 


Mean sees 
per period 


% of time 
present 


September 


1 


40 


16.2 


0.7 


58 


2.2 


47 


10 




2 


54 


8.7 


1.7 


150 


7.0 


71 


7.8 




3* 


37 


4.6 


1.4 


439 


38.8 


46 


2.6 




4 


31 


5.9 


1.1 


197 


6.2 


50 


2.5 




19 


40 


8.1 


0.8 


195 


3.5 


47 


3.2 




20 


34 


7.5 


0.8 


80 


3.1 


63 


3.3 


Total/mean/% 




38 


7.5 


1.1 


237 (140**) 


9.7 (5.8**) 


54 


4 


October 


1 


38 


8.4 


1.2 


69 


3.4 


66 


7.7 




2 


33 


6.7 


1.8 


77 


2.1 


71 


7.3 




3* 


39 


5.4 


2.9 


227 


30.0 


74 


8.5 




4 


35 


4.7 


1.2 


77 


2.9 


50 


7.3 




19 


33 


5.4 


1.0 


66 


1.9 


67 


7.9 




20 


36 


4.6 


1.4 


138 


8.0 


52 


5.1 


Total/mean/% 




36 


5.9 


1.6 


144 (87**) 


7.9 (5.0**) 


28 


7.4 


November 


1 


34 


4.8 


1.1 


139 


5.8 


80 


6.0 




2 


31 


5.6 


0.5 


142 


2.8 


64 


5.6 




3 * 


28 


5.2 


1.1 


195 


37.1 


51 


6.0 




4 


30 


5.5 


0.4 


79 


2.6 


43 


5.6 




19 


31 


2.5 


1.5 


121 


4.0 


37 


4.0 




20 


36 


7.0 


0.3 


94 


3.7 


87 


11.3 


Total/mean/% 




31 


5.0 


0.8 


154(112**) 


8.5 (5.4**) 


56 


6.1 






Displacement chases 


Dis 


3lay 


Silence 


Total time 
present 




Months 


Bower site 
number 


Mean sees 
per chase 


% of time 
present 


Mean sees 
per display 


% of time 
present 


% of time 
present 




September 


1 


30 


0.3 


23 


0.4 


70.2 


178 






2 








36 


0.6 


74.2 


396 






3* 


22 


0.3 


33 


0.1 


52.2 


472 






4 


66 


0.9 


56 


0.3 


83.5 


737 






19 


23 


0.2 


62 


0.5 


76.7 


646 






20 


31 


0.8 


49 


0.8 


83.8 


693 




Total/mean/% 




37 


0.5 


48 


0.5 


76.8 


3122 




October 


1 


36 


1.1 


59 


1.3 


76.9 


547 






2 


9 


0.2 


53 


1.3 


80.6 


539 






3* 


19 


0.6 


75 


1.4 


51.2 


554 






4 


47 


1.4 


61 


0.9 


81.6 


656 






19 


38 


0.2 


58 


1 


82.6 


597 






20 


32 


0.7 


48 


1.5 


78.7 


429 




Total/mean/% 




30 


0.7 


58 


1.2 


75.3 


3320 




November 


1 


29 


0.7 


71 


3.6 


78 


200 






2 


18 


0.4 


63 


3.4 


81.7 


250 






3* 





0.0 


44 


1.6 


49 


184 






4 


31 


0.5 


74 


2.9 


82.5 


302 






19 


10 


0.3 


84 


1.4 


86.3 


201 






20 


39 


4.0 








73.7 


128 




Total/mean/% 




28 


0.8 


67 


2.4 


76.4 


1265 





ATTENl>/VNC e AM) yiillAVIOUK BY OULJJLN l30WtHB|Rp£? 329 



Golden Boweibittis n^mfclecrf m()te thArt One 
C<il\ ol'ii given bird species. Forcxampic, both the 
^^llistic call and aiiuiii flock note of C'nmson 
Roscllas, the whistled single noU- and liie 
'chcep-checp' greeting calls ol' Grey-headed 
Robins, the whip-crack song and the 'chip-cHop' 
»llls{tf li4Stem Wbipbirds« the single Hick' note, 
and •any-tligher'' calls of Spotted 
Casbiftk, and holh 'cluick^ und coarse rattling 
fl^Fitalarm calls ol Touilvbilled Bowcr^irds, We 
once wiinesscd munierv of itie ^Ieh^^«4Iig-no^sc 
ofaduli male Victoria's Ritlebiids. 

Mimier>' was sometimes opporiimislic; thai is 
to say m inmicdiatc response to the call or 
^i^bUug of 4 gu)del For emipki a 

isKite-CSold^ Bowm^irdtefumed to his bower to 

findLaSftotWid Caihiid perched three mtlcrs f?0tt^ 
il andtfen immediately mimicked a cadurd call, 
even though ll\e visitor was silent. Ncighhout ing 
Ciolden Buwcrbirds often instantaneously 
ixrspond to eac^ ottKCft vocali^ftations, between 
males at bowers 19 and 20 located 1 "^Om apart. 
When one male gave a song of mak-^ ihe 

During S82 and S83 males gave medleys for 
8.7 and 5.9% o( their time present at bowers. 
tl'L-sL" \ ticalisalions averaging 177 and Ui-bsees, 

wspeciively (Table 4). The higher pcrccnu^ge of 
nw^cys fn S83 was primarily due toi-lhervo<ial 
eflforls of the iniraalure male at bower 3, in his 
fiist season oftcgulaf atiendance, During S82 he 

speni 35% oi'his bovver presence giving nicdiov s, 
averaging 278secs. in duration (Table 4). This 
percentage was bigli^iMizy^mpnih of S82 (Table 
5 V Tbi(t LB^ungtiire^.g^ sofm^ ija^eys ft<m the 
bower petctu untflce adltlt iTinl^?^ twr^ tdfal <>f 
^^^lins ). AduJthtolcs ga^'e medleys for an average 
»tf imly 5,4% of lime, al an average: of 1 I i soe 
duration, from favoured perches, By L-xcluding 
bower 3 data, results for SS2 were similar to those 
ofS83 (sceTables4-61.Tlie male at bovver 3 was 
b adult plumage in S83 and during this, his 
second year t>f regular bowci attendaiiee, gave 
fewer medley calls (Tabic 4), The new (but 
adult-plumaged) male at bovver 1'^ in S83 ga\ e a 
greater proportion of medleys, and for longer 
periods^ than ttther jnalps during that season Cscc 

During S82 males^iayemogre afternoon medleys 
than morning ones, a trend reflected not Just by 
the male a( bower 3 (see aboxe) but by oQier 
"individuals (Table ro. I^iinng SS3 sucli diurnal 
variation was rirvl apnareui, and males spent more 



1)60d-I^dd f2Q<^T8O0h ihan during iIicm: 

penodx in SJ?2 

ot 22S aod 125 ^ing pedodsrin S82 
and we cnnHmied a conspecific (u»iialiv 

fcmalc-plumaged) was near/at the bnwerolt 14% 
;md 2^% ofoccasit>ns. respecTiveiy. Medley ClUk 
were sometimes iJiven before/after displacement 

chases or displays (sec Table 7). 

Bower maiutciumct'. Bower maiuienancc 
involved ;i ni;itc placing a newly collected 
dccoratioivstick on itebowcr, flying doyvji to the 
ground betdWmer|»owei'^(jl tat«lri>vdcE]Bdlpa 
decoration, or flying if|J fl^ito' a towcr tff 
adjust/mov e a stick. It also included Brief visits to 
the bower to mspeci and/or to remove a leaf from 
the structure. Sontetinics ll^c owner Hew lo 4 
JtetiOl^' Vertical sapling (<lnLfrom lbe bavftirj 
spccffically to be abTjP to V^MpL hji^ J)«^ty 
adjusted bower decoKiffOrts*Mticks: before 
rctumineto continue bo\ver maintenanee. flyihg 
ofTto a favoured perch, or flymg direclly ouLoi 
the bower site. 

Males flying 10 iheirbowerlo maintain it were 
usually silent (77% of 6 15 such visits in S82 and 
SS3), but duTitig 141"! <iiI-,i.*t visits males c-illed 
brielly as lhe\ landed on their bovver perch. Such 
calls included, a single raulc with one or IWO 
scntejEtlifi) Ci^— LU1|, a.^nglc r^A3-'with ^uBat% 
fSllciwed'by WtffWtMcty w fn-^ 101, si 

single raftlf with a ru cuhi or srrctTi) call (n = 9), 
or jiisl one raffle, SL/ucaL mimicry oi vcviw/r^n -- 
20). Most such calls (04'V„) oceurred uben iv.ales 
were ()ol Carryjj^^ ^nylbin^t MaJlt?^ rWiWl^d 
ail6nt OKict deoomttng/rnaintaiiun{| boWirs. 

Mates reiut iiiiig io tbpir bpwer site witfi ^ 
decoration or sttek usually W«nt direclly to Ihe 
bower, to add ittotbc stnicnuffr. Most decorations 

(95%) were collected by males during their 
absence from tiie bower, having har\ested t'lem 
or stolen ihem from a neighbouring bovver. A few 
limes (5%) n^alcs did not leave Uic howcj area 
collect a decoililion, but merely Hew- lo a lice 
(often 4ut (jf iM.si^) to fpomy wittMUS3(tecics 
with a tiecwaflon. oa foitf ^^cih occasions ^ve 
eonnrmed tfic male had g-.'Oe to a 'store' to 
retrieve a decorauon. Wc also iwice s:;\v u rnide 
fly fiom his bower with a deeotatit^m to 'store' it 
in a tree crevice llireelnnes we saw tj male takcii 
decoration from his bower to place it Oh a 
favoured perch, only to stibscqucntly collect it 
and return it la his bower. Uniike bower decor- 
ations, most sticks (77- r.) were collected near 
.{ 1 i)-2flni disfiint ) llw bowen Sti<;Kstlo .^Os;iti long) 
Avere usually «pllttcti^ ane at4>intc» byia opiipic 



330 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 6. Variation in diurnal time-budgeted activities performed by individual male Golden Bowerbirds at 
bower sites during the display seasons of 1982 and 1983. * = data for bower 3 excluded; see Results. 





Advertisement song 


Other calls 


Maintenance 


Single 


Medley 


Season/time 


Month 


Mean sees 
per song 


% of time 
present 


% of time 
present 


Mean sees 
per call 


% of time 
present 


Mean sees 
per period 


% of time 
present 


1982 

0600-1200 
Total/mean/% 


September 


40 


10.2 


0.9 


129 


4.9 (3.3*) 


57 


5.4 


October 


38 


7.5 


1.4 


106 


5.0(1.5*) 


69 


8.8 


November 


32 


6.9 


0.5 


171 


6.2 (2.1*) 


56 


5.8 




38 


8.5 


1.1 


124 


5.2 (2.4*) 


62 


6.8 


1200-1800 
Total /mean/% 


September 


30 


2.5 


1.4 


398 


18.6 (5.0*) 


42 


1.3 


October 


30 


3.4 


1.8 


185 


12.1 (4.8*) 


51 


5.2 


November 


28 


2 


1.3 


142 


12.3 (4.8*) 


57 


6.5 




30 


2.8 


1.6 


236 


14.6 (4.9*) 


51 


4.0 


1983 

0600-1200 


November 


30 


6.2 


0.3 


114 


6.1 


41 


2.6 


1200-1800 


November 


30 


3.4 


0.5 


104 


5.7 


37 


3.3 






Displacement chases 


Dis 


alay 


Silence 


Total time 
present 
(mins) 





Season/time 


Month 


Mean sees 
per chase 


% of time 
present 


Mean sees 
per display 


% of time 
present 


% of time 
present 




1982 

0600-1200 
Tolal/mean/% 


September 


39 


0.7 


48 


0.6 


77.3 


2016 




October 


32 


0.8 


61 


1.3 


75.2 


1999 




November 


37 


0.6 


62 


2.1 


77.9 


785 






35 


0.7 


58 


l.l 


76.6 


4800 




1200-1800 
Total/meaii/% 


September 


13 


0.1 


31 


0.3 


75.8 


1106 




October 


26 


0.7 


53 


1.1 


75.7 


1321 




November 


24 


1 


73 


2.8 


74.1 


480 






25 


0.5 


56 


l.l 


75.4 


2907 




1983 

0600-1200 


November 


40 


1.0 


45 


1.2 


82.6 


2316 




1200-1800 


November 


32 


0.5 


41 


1.0 


85.6 


1536 





of times two or three sticks fused together were 
carried in. Most sticks were taken from the 
ground, but three times a male broke a dead stick 
off a sapling. 

During S82 males visited bowers 451 times to 
maintain them, adding a new decoration on 222 
(49%) occasions, and a stick on 47; remaining 
visits involving only maintenance. Of 222 decor- 
ations, 56% were beard lichen (Usnea sp.), 32% 
the creamy-white persistent flowers, or seed pods 
of Melicope (Melicope broadbentiaua), 3% 
jasmine {Jasminium kajewskii) or orchid {Den- 
drobiiim sp.) flowers, and 9% unidentified. 
During S83 males visited their bowers 1 68 times 
to maintain them; adding a new decoration on 64 
(38%), and a stick on five, occasions. Remaining 
visits involved only maintenance. Of 64 decor- 
ations, 36% were beard lichen, 20% Melicope, 
22% jasmine or Brown Silky Oak {Darlingia 



darlingiana) flowers, and 22% unseen or 
unidentified. 

Males spent more time in bower maintenance 
during S82 (5.8%) than during S83 (2.8%), and 
for longer durations per period (Table 4). There 
was no significant difference between the 
proportion of time various individual males spent 
maintaining their bowers each season (82: x- ^ 
1.33, P>0.90; 83: x- = 6.5, P>0.20), but differ- 
ences between them for the two seasons were 
significant (arcsin transformation t,o = 6.92, 
P<0.001). In September S82 most individuals 
spent less time maintaining bowers, particularly 
in the afternoon, than during October and 
November (Tables 5, 6). 

In S82 males performed 48, 22, 13 and 17% of 
total maintenance visits (n = 451) during 
0600-0900, 0900-1200, 1200-1500 and 1500- 
1800h, respectively. Most decorations (70% of 



BOWER AITEND/VNCE AND BEHAVIOUR BY (KJLDKN B(J\Vi RBIRDS 331 



fll«t mmi^ to Wi200h>. to S&3 maliwr pter- 
fbnncd 24, 2^, 30 and 1 Y% bPlottl trrnutfefianfc* 

visits III ^ \M) during '".he same periods, 
i\:spci:ti\cly. Nh^sl decorations (67% ol 64) were 
adUed in the moming (OhOO'llOOh). Of the five 
S83 slick i^ollccLion^ (^ee abovcj, (wo we iii the 

bisplacemen/ chases. When a conspecific or other 
bit^ bailed C^n d^c bower perch, or one near it. 
b&wor-cJwnmriisJiially responded flying ai, 
gnci displacing, ihe visitor to chase it out of the 
"bower site. While chasing, the bovver-owner 
ollcn fiinned his tail aiul, whefi laadiiiL! on the 
bower perch lo di-splace the visilur> giive n siniilc 
rurt/c with squeal{s). During tlic ensuing chase, 
isnsiX^ i^o: ii\clud(4 £1 sXh$1c mUis followed hy 
cka^ iiifl?or ■othcjf brieflriisnicry. 

or '95 dTsplacxiuciii chases ubsencd during 
S82, 80 were directed aiconspccifics. 1 1 al olh*;r 
birds and recipients of four were unseen. Of 49 
displacement chases seen during S83, 29 were 
directed at conspccifics. nine <\\ ulher birds, and 
icL-ipients of i I were unseen. Thus, ol' 144 dis- 
plaeemtint chases, 7fi% were directed at 
Dompccifics and 24% at other birds* 'OttwiT 
SM^IVS ill^lude<i* Wliite-thfoatcd Tree-crccpcr, 
Cnrmohat&s feucophaeus (n ^ 1 ); Bridled 
Honeyealcrin * I ): Yellow-lhroated Semb-wren 
(n = '2); Grey-headed Robm (n = 10); Basteni 
Whtpbird (n ^ 2); Bower's Shrike-thrush (u ^ ! ); 
i^d small unidentified passennes (n = 3). Iwi ol 
IfaiBse chaseii involved dispktccmcnt tVom the 
bower perch and Ihc oUiers (e.g. Yellow-thn»a1ed 
Smib-vvten. F.astertt Whipbird, Grc*y-hcaded 
Robin) iirom within 5m of ii, An Ausinihan 
I3rush-lurkey {n = 1 ), Spotted Catbird (n ^ 2) and 
Satin Bowcrhird (n - 4) perching close to ;jnd/or 
on the bower were not chased, hut w ere scolded. 

During S82 and ^83 conspecilles were seen 
vUilitigli>wers on 1 36 and 46 occasions, respect- 
ively. Of these 182 visitations; 103 w«ltV W 
fcmale-plumagcd (unmarkcd/scx liarimovm) 
tndividmils; 25 by ru>n-bo\\er-ov\ning iniiuature 
{female-pi untagcd) males (identilied by bands 
and/or behaviour); and 54 by neighl>ouring 
(marked) k^W^!^OWllef » (T^blc 7). Wc usually 
aaw only cmsts-viirftDr afwiime, on ot nca* alMWer 
ptftch, biH four times we continncd a second 
visitor simultaneously in ihu bov»ci area- 

Fenialc-plilraage<i' -hitiividiials f^cndci 
ittiknuwn) were fUltjvc as Oity apmoaclied a 
bo^en lisually via several pet*^. When they 



-And/oi' * frozen ^ i m v- 1 1 u e and peered about, without 
palting or touching deeorations. D\xsm^*i uf IQ3 
Visits by fcmale-plumaged visitrtrs, tins 
bower-owner was present al the bow er site (lahle 
7). On XO of these 9"^^ s tsit^ ihv owner lypieally 
instantly displaced and chased oil the visitor. 
mvsUy (^1 %) rK>in the hiiwci peich. ^^ftwing 
flilltsed away, some individuals jpt 8) 
fmmtidialely cirvied hack to the bower pcrcl% 
only lo be chased olY again by ibe owner 
Consecutive displacement of tlie same \ isiting 
individual fiXMnabowerpcrvfi mvt^lved 2 (n 6), 
3 (n ^ I) Of 4 (P - I) ch:. e:. Owners bricllv 
displayed lb some ( H^Va] fe3nale^uni^gedbj[rc£s 
uhcn they first anivied, f^ut dnce th« "visitar 
reached the K( .v\er percl> it was displaced and 
chased auay. ,\rier some chases (8%) oWTiers 
rctatned to ilieii howei, soinciimes aecumpanicd 
b>* (be same visitor, lo then display (Tabic 7). Qn 
190(111^99 visits 1^ fctnalo-plumaged birds tb&y 
were not chased avray ; the owner displaying io (n 
= 6)» or ignorine them as they ])crchcd abovo{n-= 
or nn (n - 5 ) ihc bower perch. Hn^ie visilorS 
may have been lenude* 

*NOn bmver-oWHing immature niales were ^trt 

10 visit bower sites tn/e limes in the owner's 
absence iind 20 limes in the owner's presence, 
these visits averaging jSsecs (Tahle 7 ). Diinn.ii ..in 
owner's absence young males spent more time 
(mean visit duration = l()4secs) on the bower, 
adjusting decorations aiul/or slicks, than wheri an 
owner was present (mean visit duralim ^ 24 
secs'i. On 1 K visits i(» the i*w ncr's itiesence young 
males were typically insianilv displaced and 
ch^ tffFby TO&6«rt>en Wd always so from the- 
bowcrjpd^K sMiptnnti^ d irr tW0 

individuals (r - 4) rnnT^Jialelx cirale3l!liiic)t(0 
die bi)wer peieli ;!ttLi tlic :hasc. only tObectUISed 
again. ( tmsccuLne .bases vliieeied ihe sain^ 
Individual iuvolvcd2(n = 2X 4 in - Hand 6 pi— 
I ) chases. Twice m owner displayed and/or 
3n«dley calls when a young male first arrived near 
Tiis bower, but when the visitor reaclved the boww 
pL-Tcli it w.is displaced and chased. Mlcr two such 
chases ihc owner rciurned die l iowec to disphiv 
bf^(|fjy '(Tah)c 7). On two Msits in one ow-mt's 
presence* yuung m4ilps wc^rc not cl^ascd ^w^ay, put 
this was because theTnal<MTWDet 6f tovW^ W*» 
immature lie tolerated another young male on 
liis bower tor 100 and I39secs (see also beU>w>. 

Male bower owners visited adjacent msiJes^ 
bowers 54 limes, 35 being in an owner's absence 
and 19 in an owner's presence. On 1 1 of the lAltcr 
visits iheintrudcr wwclx^sf^o^llirbowe^ 



332 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 7. Conspecific visits to six Golden Bowerbird bower sites (data for S82 and S83 combined) and the 
bower owners reaction to them when in attendance. 



Visiting conspecifics 


Female- 
plumaged: sex 
unknown 


Non-bower- 
ownins voung 
males 


Bower- 
owning males 


Total/Mean/% 


Visits to bower 

sites 


No. to bower perch 


76 


22 


50 


148 


No. to other perches(<5m distant) 


27 


4 


5 


35 


Total number of visits 


103 


25 


54 


182 


Mean time per visit (sees) 


52 


38 


17 


36 


Bower-owner absent (No.) 


4 


5 


35 


44 


Bower-owner present (No.) 


99 


20 


19 


138 


Reaction of bower 
owners to visitors 


Displacement 
chases 


No. from bower perch 


64 


18 


10 


92 


No. from other perches (<5m distant) 


15 





1 


16 


Total number 


80 


18 


n 


109 


% of times owner present 


81 


90 


58 


79 


Displays 


No. before a chase 


14 


2 


1 


17 


No. after a chase 


6 


2 





8 


No. without a chase 


6 








6 


Total number 


26 


4 


1 


31 


Medley calls 


No. before a chase 


6 


1 





7 


No. after a chase 


19 


1 


1 


21 


No. before a display 


6 








6 


No. after a display 


21 








21 


No. with no chase or display 


10 


2 


1 


13 



by the owner, but 4 times successftiliy stealing a 
decoration. Displacement chases twice involved 
body contact between adult males. One owner 
rapidly displaced a rival from his bower perch to 
grapple with him, the two tumbling toward the 
ground before separating. The marauder then 
flew off, pursued by the owner. On 8 of the 19 
intrusions there was no chasing; twice the 
intruder flying off at the owner's return before it 
could steal. On 6 occasions a thief stole un- 
molested in the presence of the immature male 
owner of bower 3. 

Thieves managed to steal a decoration during 
83% of their visits to bowers of rival males, 
obviously being most successful (100%) in the 
owner's absence. Time spent at a rival's bower 
was brief (mean = 17secs). Decoration theft 
between adjacent bower-owners was rife. Having 
apparently noted a neighbouring rival male's 
absence, by lack of his calls, bower-owners often 
flew immediately in the direction of the presum- 
ably unattended bower to then immediately 
return with a decoration. For example, when the 
male at bower 19 was absent (DWF observing 
there), his immediate neighbour at bower 20 
(CBF simultaneously observing there) would fly 



to bower 19 and immediately return to his own 
bower with a stolen decoration. 

During our second (14 September) S82 ob- 
servation, the immature owner of bower 3 was 
challenged for the site by a male in his first year of 
adult plumage. Both birds were present at the site 
for 211 of the 360min observation. Some of this 
time they perched close to each other, the 
immature owner continuously giving medley 
calls with mimicry, frog-like notes and scolds 
(for 139mins); and the adult-plumaged 
challenger frog-like notes, scolds and sometimes 
mimicry. At other times they chased each other in 
prolonged tail-fanning flights about the bower, or 
in short flights from perch to perch while 
fluttering/flicking wings in agitated manner. The 
adult was mostly chased by the immature, but 
sometimes this was reversed, the immature twice 
displacing the adult from a perch but four times 
the reverse. The adult once performed Bow and 
Head nod and shake displays. The immature 
retained his site, however, and attained adult 
plumage the following season. 

Males spent little (<I%) time present at bower 
sites in performing displacement chases (Tables 
4-6). Displacement chases averaged 34 sees, but 
varied much between individuals. 



fiO>W£R ATTENDANCE AND BEHAV JOUK iS\ (jULLJi:.N bOWrrUJIKIivS 



Di^lays. Males pcribnncd three basic display 
elemenls {Bow^ Heud maJ <xnU shaken and 
FUght/hover) al bower sites, m the Sbw tfte male 

lowered his bill an-j hcaJ ngi-ity i.lo\vn\v;irds ntifl 
cr-vCtcd his ciuwi. atui nape featlicrhig. thus 
emphasising his bnliianl yellow 'cresi" vvtiilc Ins 
V' I'gs were shghlly io fuiJy drooped (depending 
ii display hllensity) imd occasiiofiiilly llickc*}, 
The tail was pulled forward bcncati] the lowered 
head* and someliTnes held to ont: ?*id6. 

The Hl'ihI nod and shake involved the male 
nodding his ngidly downw^ird-puinling hill dnd 
bead up and down with erect head plumagr and. 
svhenuisiplaying iulwseJy^aJso shakitig his head 
from side to side. As he nodded he peered ituenrly 
ar and sometimes pecked, his perch (usu:illy a 
vertical sapling trunk). This display also 
eiQphasised the^TvlIow central crown and nape 
*cr«st\ The wing^vvere slight!)' mjfulfe ^oped 
aj>d owasiandw flotttwdrflw N^ad nod 

ami shake or vice ver-^a. niuy be perlbnned 
ciinseeiuivcly on the same perch, when display mg 
isitktense. 

(n Fli^^hi 'liover disphiy a male f\c\\ with slow 
and deliberate 'butterfly- like* beais ol exicnsively- 
opencd w ings aroiund his bower at 0.5-4m ( usually 
l-2m) above ground*, oucui^ionall^ WitJi a 
decoration i»i his bill. During this display fli^ liic 
tail may he fully tanned dunng brief mid-fliglii 
hovers: cxposint: tlu- ncIIow outer rectriees. 
Also in tliis display, males typically si"opped lo 
llOVer in Iroul of a vertical sapling trunk or its 
leaveai, With bill tip touching the stcnVleavcs, wtulc 
Ttspeatedlv and alternately fiilly fanningaiid closing 
llie tail. 'Ttie bird then Hew ^butterfly-lilSfi!"* to 
.another sap hug to pertbmi tlic hover/tail fanning 
display again. During Flfghf/kover the male may 
hriePly perch on a vertical sapling stem where he? 
may also perform a Hecul nod u^/r/.v/w^-e display. 

Each display consisted ot one, two, or all the 
above three elements. An element was often 
tiepesded during-^3 display. During S82 and S8^ 
WtecOtded a tcH«f 6ri46 displays, involving 
238 separate elements: 62 Bows. 104 Head nod 
mdshakes and 72 FUghldiovtr^ (Table 8). Of the 
62 Bow elements: 48"/i> were perlbnned on bower 
perches, 44% on vertical sapling trunks- ^ud 8% 
.0ftl^ra<tfrealT»erches. The Bow waspeifeimed 
altmc 124^) or, during other displays, was 
perfbmled"beforc (23%) or after (29%) Head nod 
and shake. Or before (11%) W afef (13%) a 
FUghtdmet (see "ftblc 8). 

Of 41 (ot«l l<M Head fu}d and shakes: 13% 

were p^^bnncd on ba^wt pmlmi 75% m 



vertical sapUng frUtlksi «iml 12% on hori/.onttd 
perches (TaM^ Most (90%) times a m^k 
performs^ tbo ilem rim/ and shiiltt ixit Onepmli 

only, hut a tcv. tioics ne\v!ctnne(n-5)ortVHt(ll 
^ 5) addiilon^J perches in eontimjous display. 
I k //.zL-t/ .'i^>j urn! shiike was performed alone 
(3S/b) or, during oQier displays, was perfurmwl 
Utbrc ot after (14%) Bow. tir bclorc 

{I4%)orafta:n7%) flight/ hoi evisa^ hbk ^) 

Of ^ total of 72 Flli^hf/himr displays: 96% 
were performed al vertidEd sapling trunks Of 

leaves and tjie lemainder at decorated howw 
platlorms. prior :c landing on the buwer p^ffch. 
The number itf hovers L!i\en during a 
F/ ighi/ hover dtspUiy viicied. Mosl iimcN^n=52)d 
niaic performed only one luvvei; but at Other 
times pcrtgfiK>e4 2 (n = 9), 3 (n =5), 4in=] Joe 5 
{n = I), Kour flight displays Isdt^ f liwur, 
Flij^ht/hovcr was performed alone .t. 
durmg other di'>[il;iys. vsas pcTtbiincd helbie 
( 11%) or alter 1 Bow, or hctbre (25%) or 
after (21%) Ucoi^ nod and shake (sec Table i«>. 
We Ci">uld disecm no clear sequence in which tfiie 
three display elements might typically be per- 
Ibmicd, possibly because no complete succe&sHd 
courtship, to CopuiMtoilt WOS^b^W^ 

"VrTien n'iales landed on their bower percliio 
di^play {Bow or Head noij and shake) they 
someiinies gave oije- ratde and stntciil but i>u 
other perches gave only a scjueaiyn). Dining 
Fiighf/Hover displays males often gave a 
sqNeal{s) and-or hncf nvmicry. iis Hying iVoiU 
one perch to iinni -icr 1 1'-i "!e S]. Dunng Bow and 
J lead nod ana' shiike L^ivcn -ri bi'ncr aj'd 

other perches, males occasionally held a 
decoration in the bill, but only twice when a 
femole-piiiinagi^ individual ^\^iuu>.wn to )>e in 
theittvd. JMtit$t)r$ap1ing?iised&imigdisp3aiys 
averaged 2.1 ^ 1.3m above groundlmi4wer*35 
± \.Hm (n - 36) from bowers. 

Total numbers ol elements performed durii^g 
eacti d^sp^ay ttneludM\g rcfietltion ol" any 
elements; see above) are summarised in Table 9. 
Mesl displays involved one (63%), two (23%)or 
thrcc4i6%) plemcot^ bulfiv« di^flays consisted 
dr^t^^:0l^3Yand rune. Of ehe 238 «teni«Dte 
ob$er^: IB X'iA per disphiy) \yac peifonnici 
exchisiViily on hower jK-rchcs, 137 (1.4 per' 
display) nr.'at verliciil perches, 13 (1,1 per 
display) ut] bPTi2oi\tal j[>erc;lTcs and 70 (3.0 per 
displ^) on a^ottnWnaliwi of wich^pcs (Tabic 
9 1 There was no apparent S^oepce 10 4i^lfty 
clemenls- Of total dl<5|>1ay inglfnic 'onty l^WIB 
Qifdie'bowerpprfih. 



334 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 8. Number and sequence of display elements (n = 238) performed by male Golden Bowerbirds at their 
bower sites during 146 displays, and the number of times males called and carried a decoration in their bill 
during display (data for display seasons of 1 982 and 1983 combined). * = hovers performed in front of vertical 
trunk or its leaves; ** = hovers directed at the bower tower before landing on the bower perch; see Results. 



Display elements 


Bow 


Head nod and 
sh^kc 


Flight/ 


Total number of 


% of total 


Display 
elements 


Number given 
on/directed at * 


bower perch 


30 


13 


3 ** 


46 


19.3 


vertical support 


37 


79 


69 


175 


73.6 


horizontal perch 


5 


12 





17 


7.1 


Total number 


62 


104 


72 


238 




Number of single elements only 


15 


39 


24 


78 


32.8 


Number given 
with other ele- 
ments 


before a bow 




18 


8 


26 


10.9 


before a nod& shake 


14 


. — 


18 


32 


13.5 


before a flight/ hover 


7 


15 




'*2 


9.2 


after a bow 




14 


7 


21 


8.8 


after a nod& shake 


18 




15 


33 


13.9 


after a flight/ hover 


7 


15 




26 


10.9 


Total number 


62 


104 


72 


238 




Calls 


No. given on bower perch 


12 


8 





20 


8.4 


No. given on other perches 


4 


13 


14 


31 


13.0 


Decoration 
held in bill 


No. held on bower perch 


10 


4 





13 


5.5 


No. held on other perches 


3 


6 





9 1 3.8 



Numbers of elements during a display did not 
increase as S82 progressed, possibly because it 
was extremely dr>'. During September, October 
and November 56, 46 and 61% of displays 
involved one element, 33, 29 and 26% two, and 
11,11 and 18% three elements respectively. Two 
six-element displays occurred in late October, 
and one four-, one six- and one nine-element 
display in November of S82. During November 
of S83, display involved one (79%), two ( 1 9%) or 
three (2%) elements. 

Of the total 146 displays by bower-owning 
males we saw a female-plumaged individual 
simultaneously at the bower site 26 times (22 on 
the bower perch and 4 perched close by; see Table 
7). On these 26 occasions males displayed 
before/after a displacement chase (n = 20) or 
displayed only (n = 6); performing one display 
element (n = 17: five Bows, five Head nods and 
shakes, and seven Flight/hovers), or two (n = 5), 
three (n = 3) or nine (n = I) elements. 

The longest display observed (I7mins) 
involved performing display elements nine and 
three times, each group of them interspersed with 
much medley calling. A female-plumaged 
individual was perched on a vertical sapling 
initially, but then moved to the lower side of a 
bower tower to perch motionless, with sleeked 
plumage, and stare at the male. Once, when the 
visitor landed on the bower perch, it was 



immediately displaced and chased by the owner. 
As in all other display/calling sequences with a 
female-plumaged individual present, this did not 
terminate in copulation. 

Males spent little (1.1%) time displaying 
during S82 or S83 (Table 4). During S82 fewer 
and briefer displays were recorded in September, 
during both mornings and afternoons, than in 
November despite decreasing bower attendance 
during the latter (Tables 5, 6), Mean display 
duration was 67 sees (Tables 4-6). 

Silence and other behaviour. Males spend much 
time silently on perches above their bower, when 
they preen, bill-wipe, sun, change perches, turn 
180° to face the opposite direction to sing or to 
better listen to neighbours' calls, or forage 
locally. Wlien males returned from an absence, 
having obviously bathed, they flicked their 
wings, shook and fluffed their damp feathers and 
continuously preened. Once during a brief rain 
shower a male shook his wings and fluffed his 
plumage, before bathing by flying into sapling 
foliage and briefly fluttering/hovering amongst 
the wet leaves. On two occasions (at 0942 and 
1245) different males perched in direct sunlight 
above the bower and sunned themselves; with 
erected breast, rump, head and nape feathers, 
down-pressed tail and drooped wings. We 
witnessed males fly from a favoured perch (n = 
28) to snatch an insect from nearby foliage or 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 335 



tnmk, or to hawk (n - 2) an insect from the air. 
Males were twice seen to fly to take a vine fruit, 
tw ice to feed on a fruit on the ground and twice lo 
retrieve fruit Irom a food store near the bower (a 
crevice in a vertical trunk). 

The proportion of time at a bower that males 
spent silently (other than during bower main- 
tenance periods, displays or chases) averaged 
76% during S82 and 84% during S83 (Table 4). 
The lower S82 figure was in part due to the 
continuous medley calls given by the male 
establishing himself at bower 3. Periods of 
silence were similar for each individual for each 
month (Table 5), and during different times of the 
day during S82 (Table 6). 

DISCUSSION 

ANNUAL SEASONALITY AND RELATIVE 
LEVELS OF BOWER ATTENDANCE BY 
MALES. Paluma Range male Golden Bower- 
birds typically started attending traditional 
bowers in late August/early September, as 
temperatures initially rise from midwinter ones, 
approximately 6 to 8 weeks before females 
commenced egg-laying (Frith & Frith, 1 998). On 
the Atherton Tableland, especially at slightly 
lower and thus wanner altitudes, the display 
season starts a few weeks earlier (Marshall, 1 954; 
Frith & Frith, unpubL data). Bower attendance 
declined in December; particularly when pre-wel 
season rains commenced (Warham, 1962; Frith 
& Frith, 2000b and this study). During the wet 
season proper, few advertisement songs were 
heard, decorations deteriorated, and few males 
briefly visited bowers lo add decoration during 
dry spells. Males moult at this time (Frith & Frith, 
unpubl. data). A brief period of activity occurred 
in late March-early May, when bowers were 
poorly decorated and few songs given, as noted 
by Warham (1962) on the Atherton Tableland. 
This post-courtship activity* is in part reflected by 
infrequent attendance of traditional bowers by 
immature males, a situation also found in 
Tooth-billed Bowerbirds at the same location 
(Frith & Frith, 1994; 2000b). 

Seasonal variation in time invested at/on 
bowers by male Golden Bowerbirds may vary 
year to year subject to prevailing weather 
conditions, particularly excessive wet season 
rains and drought, and thus food resource 
availability (see Lenz, 1993; Frith & Frith, 1994; 
and this study). During abnonnally dry seasonal 
conditions (as in S79 and S82 of this study), when 
rainforest fruit crop was poor and invertebrate 
numbers and biomass low, not only was the 



display season shorter and male attendance at 
bowers reduced, but fewer females attempted to 
nesl or did so successililly (Frith & Frith, 1998). 
A similar situation was recorded for Paluma 
Range Tooth-billed Bowerbirds during the dry 
S79 (Frith & Frith, 1994; 2000b). During season- 
ally typical conditions, male Golden Bowerbirds 
attended their bowers at consistent levels 
throughout the day, as did Macgregor's 
Bowerbirds Atyihlyornis macgregoriae (see 
Pruett-Jones & Pruett-Jones," 19^82), but in 
adversely dry conditions they did so almost twice 
as much during the mornings than during 
afternoons. Drought conditions had similar 
impacts upon both sexes of a polygynous, 
lekking, neotropical hummingbird (Stiles 1992). 

Adult male Golden Bowerbirds (n = 7 ) spent an 
average of 50% (range 32-69%) of daylight 
within 15-20m of their bowers, at a mean of 2.9 
(range 2.2-4.2) bower visits per hour and each 
averaging 9.5mins in duration. The former 
figures are similar to those found for male 
Macgregor's Bowerbirds (n = 5) that spent an 
average of 54% (range 20-75%)) of daylight 
within 15-20m of their bowers, but did so at a 
mean of 1 .4 bower visits per hour (range 0.6-2.0) 
and each averaging 4.6mins in duration (Pruett- 
Jones & Pruett-Jones, 1982). Both species are 
predominantly frugivorous (Pruett-Jones & 
PrueU-Jones, 1985; Frith & Frith, unpubl, data). 
Male Tooth-billed Bowerbirds, almost ex- 
clusively frugivorous during their peak com"tship 
and mating season, spent an average of 64%> of 
daytime at or near (< 1 m) their courts, at a mean 
of 2.9 court visits per hour and each visit 
averaging 23mins in duration (Frith & Frith, 
1994). Male rainforest Satin Bowerbirds, with a 
67% fruit component of annual diet, spent an 
average of 73% of daytime within 50m of bowers 
(Donaghey. 1981). A male Great Bowerbird, 
ChJamydera nuchalis, a species considered 
predominantly frugivorous (Diamond, 1986a; 
Schodde & Tidemann, 1988), but probably less 
so than the above species, spent 47% of daylight 
hours at or near his bower at peak mating season 
(Veselovsky, 1978). 

Regent Bowerbird, Sericulus chrysocephalus, 
annual diet includes 81% fruit, and yet males 
spent a mere 3% of daytime at or near their 
bowers at an average of 0.43 bower visits per 
hour, and each visit averaging 6.7mins in 
duration (Lenz, 1994). Similarly, a Flame 
Bowerbird, S. aureus, spent 6% of daytime 
at/near its bower (Dwyer & Minnegal in Coaies, 
1990) and a Fire-maned Bowerbird, S. bakerU 



336 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 9. The number of display elements (n = 238) performed by male Golden Bowerbirds during 1 46 displays 
on the bower perch and elsewhere in the bower site (data for the display seasons of 1982 and 1983 combined). * 
= these include bower perches, horizontal perches and vertical sapling trunks that Bow and Head nod and shake 
display elements were performed on, and vertical tree trunks or its leaves that the Flight/hover display were 
directed at; ** = during a display any element may be performed more than once; see Results. 







Number of display elements per display ** 


Total no. of 

displays 
(% of total) 


Total no. of 






one 


two 


three 


four 


six 


nine 


display 
elements 
{% of total) 




Bower 


12 











1 





13 (8.9) 


18(7.6) 




Vertical 


68 


21 


9 











99 (67.8) 


138 (58.0) 


« 


Horizontal 


10 


1 














11(7.5) 


12 (5.0) 


c 
_o 

"S 

o 


Bower to vertical 





7 


1 











8(5.5) 


17(7.1) 


Bower to vertical to bower 





1 


1 





1 





3(2.1) 


11 (4.6) 




Vertical to bower 





2 


1 











3 (2.1) 


7(3.0) 




Vertical to bower to vertical 








2 


I 


1 





4 (2.7) 


16(6.7) 




Vertical to horizontal 





-1 


2 








1 


5 (3.4) 


19(8.0) 


Total number 
(% of total) 


91 

(63.1) 


34 
(23.2) 


16 

(10.2) 


1 

(0.7) 


3 

(2.1) 


1 

(0.7) 


146 


238 



<1% of daytime at its bower (Mackay, 1989 and 
in Lenz, 1 993). An explanation poslTilated for this 
exception is that regent bowerbirds, Sericulus 
spp., represent an early stage in the evolution of 
bower-building, in which bowers have not yet 
replaced elaborate male nuptial plumage. Male 
Regent Bowerbirds initiate courtship in the forest 
canopy, before accompanying the female to the 
bower where a prolonged courtship display 
primarily presents colourful nuptial plumage 
(and less so bower/decorations) to the female 
(Lenz, 1994). Thus the bower plays a less sig- 
nificant role in courtship. 

It has been observed that a disproportionately 
large percentage of tropical rainforest-dwelling 
passerines, with a polygynous mating system 
based upon court/bower/arena displaying pro- 
miscuous males, are predominantly frugivorous. 
The seasonal abundance of rainforest fruits, 
economically undefendable because of their 
spatial/temporal distribution, both promotes the 
emancipation of males from nest duties and 
enables females to raise offspring unaided by 
conspecifics(Snow, 1976, 1982; Frith &Beehler, 
1998). This said, Donaghey (1981) found that 
both adults and nestlings of the monogamous 
Green Catbird, ^/7z/wc^^/a/,v crassirostris, are more 
frugivorous than the polygynously breeding 
Satin Bowerbird, and noted that frugivory is but 
one of many factors involved in the evolution of 
avian promiscuity, Male Golden Bowerbirds also 
store, or cache, fruit foods around their bower site 
(Frith, 1989 & pers. obs.), as do male 
Macgregor's Bowerbirds (Pruett-Jones & 



Pruett-Jones, 1 985). Such storing of fruits around 
bowers would enable males to spend inore time in 
bower attendance. 

MALE BEHAVIOUR AT BOWERS. Habitual 
perches, vocalisations and silence. Bower- 
attending male bowerbirds studied to date 
advertise their bower location with specific calls 
given (Tooth-billed Bowerbird excepted) 
relatively infi-equently from favoured perches 
(Gilliard, 1969; Veselovsky, 1978; Donaghey, 
1981 ; Frith & Frith, unpubl. data). Male Golden 
Bowerbirds gave the distinctive bower advertise- 
ment rattle^ single notes [squeal, screechy 
scold-rasp and wolf-whistle) and a medley of 
calls that included much mimicry of frog- and 
cicada-like notes, and fine vocal avian mimicry, 
from habitual perches. The Tooth-billed, 
Archbold's (Archholdia papiiensis), gardener 
{Amblyornis spp.), Regent, Satin, Spotted (Chla- 
mydera maculata). Western (C. guttata). Great, 
and Fawn-breasted (C. cerviniventris) Bower- 
birds include avian mimicry in their bower 
advertisement and/or other non courtship 
vocalisations (Marshall, 1950; Gilliard, 1969; 
Bradley, 1987; Lotfreddo & Borgia, 1986; Frith, 
1989; Lenz, 1993; Frith & McGuire, 1996; Frith 
ct ai.. 1996, & pers. obs.). Male bowerbirds 
usually use a distinctly different, quieter and 
more complex, subsong in courtship display that, 
in the case of all of the above except the 
Chlamydera species, includes avian mimicry. 
During this study we did not hear Golden 
Bowerbirds giving subsong mimicry while 
displaying, but have done so subsequently. We 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 337 



observed thai long medleys of calls including 
much mimicry were given before/after display 
posturing, and particularly if a female-plumaged 
bird was present (see Table 7). We think it hkely 
that this is more informative to females than 
advertisement song, as it has been demonstrated 
that female Satin Bowerbirds use the quality of 
more intubate male mimicry to assess the relative 
merits of prospective mates (Loftredo & Borgia, 
1986). 

Immature males lacking a bower gave medley 
calls with mimicry when visiting bowers, as did 
younger adult males in their first year of bower 
ownership (Frith & Frith, 2000b and this study). 
Adult males gave fewer medleys with mimicry, 
but it is possible that the quality of their avian 
vocal mimicr}' is higher than that of younger 
birds. Among competing male Satin Bowerbirds, 
older males produce longer bouts of higher- 
quality avian vocal mimicry than do younger 
males and also gain higher matins success 
(Lofiredo & Borgia, 1986). Thorpe (1985) stated 
that there is some evidence that variety in male 
bird song is attractive to females, and suggested 
that mimicry may simply be a way of increasing 
repertoire size. Robinson & Curtis (1996) 
demonstrated that most mimicry content of lyre- 
bird (Menura spp.) calls is learned, is culturally 
transmitted, and its quality and sequence (of 
model spp) could therefore provide conspecifics, 
particularly females assessing male quality; with 
a clear indication of potential mate experience/ 
age/survival. 

Male Golden Bowerbirds spent 80% of their 
time present at bower sites in perching silently 
above the bower, this being 9% more than in 
Macgregor's Bowerbird (Pruett-Jones &, 
Pruett-Jones, 1982). Wliile adult male Regent 
Bowerbirds spent only 1 7% of time present at the 
bower site in perching silently this figure merely 
rellects the small proportion (3%) of total 
daylight they spent at bowers (see above; Lenz, 
1994). It would appear that male Satin Bower- 
birds spent 87% of time present al the bower site 
in silence (Donaghey, 1981: 181-182). In marked 
contrast, male Tooth-billed Bowerbirds spend 
<2% of time perched at the court in silence, most 
of their time there (96%) being spent smging 
loudly. Court advertisement vocalisations of 
Tooth-bills are thus much more frequent, males 
having no epigamic adult plumage but a most 
elaborate vocaT display (Frith & Frith, 1994). 

Bower mainlencmce. Male Golden Bowerbirds 
spent an overall average of 4% of time at the 
bower site in maintaining the bower structure/ 



decorations. In more typical climatic conditions 
males spent an average of 3% of time at the bower 
site in bower maintenance, but during adversely 
dry conditions spent almost twice this time doing 
so. The limited time birds spent on the bower 
presumably reflects (a) the low maintenance 
required once it is largely built (given its llingus- 
tused and 'traditionarnalwe) and decorated; and 
(b) the fact that adult male nuptial plumage 
remains a predominant part of courtship in this 
species. Limited compatible figures for bower 
maintenance by: Macgresor's (12%; Pruett- 
Jones & Pruett-Jones, 1982^), Tooth-bills (1.2%; 
Frith & Frith, 1994), Regent (61%; Lenz, 1994), 
Satin Bowerbirds (8%> of all daylight; Donaghey, 
1981) are variable. The figure for Tooth-bills is so 
low because males spend so much of daylight 
above the court (but at its site); whereas the high 
figure for the Regent, which builds a most 
nidimentaiy and sparsely-decorated bower, is 
because males spend little time at the bower site 
(Lenz, 1994). 

In restricting its decorations to beard lichen, 
melicope seed pods and whitish flowers, the 
Golden Bowerbird is far less catholic in bower 
decorations than all other polyg\'nous bower- 
birds with the exception of the Tooth-bill which 
uses only leaves of various plants (Frith & Frith, 
1993, 1994) and the Fawn-breasted Bowerbird 
which uses only green fruits, leaves and the 
occasional flower (Peckover, 1970; Pruett-Jones 
& Pruett-Jones, 1994). Bower decorations of 
greater significance to some bowerbird species 
are items rare in the birds' environment (Frith 8c 
Frith, 1990c; Frith etal., 1996), and an abundance 
of such decorations on bowers enhances the 
matins success of the bower owner (Boraia, 
1985b^ 1986; Borgia & Gore, 1986). Thus, rare 
decorations might indicate something significant 
to females, and rival males, about the owner's 
fitness/dominance levels. But are bower decor- 
ations used by Golden Bowerbirds rare in their 
habitat? The answer needs to be framed in the 
context of extensive undisturbed upland rain- 
forest, lacking the roads, tracks and clearings of 
today. In this context, we suggest that melicope 
seed pods wmild have been relatively hard to 
find, as M. broadhentiana is a pioneer shrub 
(Hyland & Whiflln, 1993) that is today found on 
track/clearing edges. Before the latter were 
available the plant would have been largely 
confined to areas of small-scale natural forest 
damage, such as larger tree falls and cyclones. 

While beard lichen is far more widespread in 
upland forest than the melicope it is sun-loving 



338 



MEMOIRS OF THE QUEENSLAND MUSEUM 



and would, in extensive primary upland forest, 
have been predominantly confined to woody 
twigs and branches of upper canopy and 
emergent trees — an exposed part of the forest 
not typically frequented by Golden Bowerbirds 
(pers. obs.). Fresh orchid flowers are never 
spatio-temporally abundant in Australian upland 
rainforest and in any event, like the whitish 
tlowers of other plants, provide inferior bower 
decorations because they will and need replace- 
ment. Golden Bowerbirds can thus be seen to tit 
the broad pattern of male bowerbirds using some 
items that are relatively rare as bower 
decorations. 

Aduh males brielly visited the bower of rival 
males, usually in their absence, in order to steal 
bower decorations. Such decoration theft has 
been documented for several other bowerbirds; 
includina the Tooth-billed (Frith & Frith, 1993, 
1994), Voeelkop (Diamond, I986a,b, 1987, 
1988), Regent (Lenz, 1 994), Satin (Borgia 1985b; 
Borgia & Gore 1986; Hunter & Dwyer, 1997), 
Fawn-breasted (Coates, 1 990) and Yellow-breasted 
(C. lauterbachi) Bowerbirds (Pruett-Jones & 
Pruett- Jones, 1994). 

Bower marauding is known in Macgregor's, 
Vogelkop, Regent, Satin, Spotted, Fawn-breasted 
and Yellow -breasted Bowerbirds ( Pruett-Jones & 
Pruett-Jones, 1994). While male Golden Bower- 
birds may (but not observed) steal the odd, 
unfused (i.e. recently placed), stick from the 
bower apex of a rival male, we did not see any 
attempt to damage ('maraud' of Pruett-Jones & 
Pruett-Jones 1994: 609) a bower of a rival. Bower- 
owning male Macgregor 's Bowerbirds attempt to 
damage bowers of rivals as well as steal their 
decorations, including the moss of the lower base 
(Bulmer in Gilliard, r969: 305; Pruett-Jones and 
Pruett-Jones, 1982; pers. obs.). Stealing of the 
latter is noteworthy, suggesting it may function as 
decoration (analogous to beard lichen on Golden 
Bowerbird bowers) and not a structural element. 

It has been demonstrated that a strategy of 
bower decoration theft hy males is an evolution- 
ally stable one, as opposed to the contrary 
strategy of guarding bowers and not stealing 
(Pruett-Jones & Pruett-Jones, 1994). Male Satin 
Bowerbirds with more decorations on bowers 
tend to steal more often than they are stolen from 
(Borgia & Gore, 1986). This is because relative 
levels of bower decoration enable females to 
assess an individual male's quality (fitness), 
based upon his success in conflict with rival 
males. The greater numbers of more favoured 



decorations on a bower positively inlluenced 
relative male mating success (Borgia, 1985a,b, 
1986; Borgia et af, 1985; Pruett-Jones & 
Pruett-Jones, 1994). Bower quality has also been 
found to correlate well with relative male mating 
success in both Satin and Regent Bowerbirds; 
and males of both species mostly maraud and 
damage bowers of their nearest neighbours, their 
most likely sexual competitors (Borgia, 1986; 
Lenz, 1 993 ). The reason for theft by rival males is 
thought to be sexual selection resulting from 
females choosing to mate only with males 'hon- 
estly advertising' their fitness with such 'rare' 
bower decorations (cf Zahavi & Zahavi. 1997). 

Displacement chases and displays. As 8 1 % o f al I 
female-plumaged conspecitlcs perching on the 
bower were immediately displaced and ag- 
gressively pursued out of the bower site by the 
bower-owning male Golden Bowerbirds it is 
likely, in view of what is known of other 
bowerbirds, that such behaviour typically greets 
females as well as adult and immature males. Of 
the 1 9% of visiting female-plumaged birds not 
immediately chased otf, half were displayed to 
and half ignored. Adult males displaced and 
chased immature and adult males from their 
bower/site but we only twice saw physical 
combat, as did Chisholm & Chaffer (1956). 

Display by a male concealing himself from a 
visiting female, by crouching behind a court tree 
or central maypole bower base to give subsong 
with mimicry, is typical initial Tooth-billed, 
Macgregor's and Streaked {Amblyornis 
siibalaris) Bowerbird courtship (Diczbalis, 
1968; Gilliard, 1969; Frith & Frith, 1993 & pers. 
obs.). We saw no male Golden Bowerbird 
attempting to hide from a visiting female to give 
subsong with mimicr>' during this study, but have 
subsequently done so. In hindsight, we 
understand this was due to limited field of view 
from hides, a point of great importance to 
students of bowerbird behaviour (Frith & Frith, 
unpubl. data). 

It is probable that the three basic display 
elements we obser\ed are performed during 
successlul courtship, perhaps in a typical 
progressive sequence, but as we witnessed few 
displays to (unsexed) female-plumaged birds (n 
= 26), and no copulations, we could not confirm 
this. The Head nod and shake display is only 
broadly similar to postures and movements 
known to be performed by courting male 
Gardener and Archbold's bowerbirds (Gilliard, 
1969; Frith et al., 1996; pers. obs.). The Bow 
display, which enhances the contrastingly 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



339 



brillianl yellow mid-crown patch and nape 
'crest', appears unrecorded in other bowerbirds. 

The Flight/hover display is unique to the 
Golden Rowerbird. This is not surprising, as this 
display clearly fLmctions to visually present both 
the (uniquely within Ptilonorhynchidae) brilli- 
antly coloured dorsal and ventral contour and 
night plumage of adult males. The deliberate 
slow flight display punctuated with hover(s) with 
conspicuously repeated tail-fanning, to expose 
the pure yellow outer rectrices, is visually 
spectacular. This might be perfonned with a 
bower decoration held in the bill (ChatTer in 
Chisholm & Chaffer, 1956; and this study). The 
closest any other bowerbird comes to a courtship 
night display is the vigorously repeated 
to-and-fro 'extra-bower' fluttering flight/leaps, 
between vertical sapling stems, by closely related 
Macgregor's Bowerbird (Stevens in Greenway, 
1 935; Mackay & Cheeseman, 1 990; pers. obs.). It 
has been noted that bowers of Macgregor's 
Bowerbird are often built adjacent to numbers of 
vertical sapling trunks (Gilliard, 1969: 302; 
Pruett-Jones and Pruett- Jones, 1982), and they 
might be a prerequisite bower site feature to 
accommodate the ' flight' display. In the light of 
this, and in view of the elements of its Flight/ 
hover display, it is possible saplings appropriate 
for hovering at/perching on might influence 
bower site selection by Golden Bowerbirds. 

Male Golden Bowerbird behaviour at bower 
sites is mostly cryptic, given they are displace- 
ment chasing and/or displaying for <2% of their 
total time present there. Males apparently 
depends largely upon bower/decorations and, 
subsequently, their colourful plumage to impress 
females, rather than a complex bower. Thus, il 
has been obser\'ed that in this bowerbird, unlike 
most, untidy bower construction and variation in 
their shapey'bulk suggests gross bower features 
are of less significance to females than is the 
discrete and relative small part of them modified 
into a 'platfonn(s)' for the exclusive placement of 
decorations (Frith & Frith, 2000a). Maintenance 
and decoration of the platfonn(s) requires but a 
small proportion of bower attendance time, once 
the basic bower is accumulated. The platfonn(s) 
does, however, provide a quickly and easily 
located 'marker' (cf Borgia, 1985a; Borgia el al., 
1985) for females seeking older, more exper- 
ienced, males to assess as potential mates. 

No data were obtained on relative reproductive 
success rates within male Golden Bowerbird 
populations. The possibility that older males are 



more successful than younger rivals has been 
found, or indicated, to be the case in other 
non-lekking bowerbirds (Borgia, 1985a) and in 
unrelated lekking passerines in which pro- 
miscuous males court females at traditional sites 
(Lill, 1974a,b; McDonald, 1989a,b, Andersson, 
1991 ). Clearly, promiscuous adult males estab- 
lishing themselves within a lek, exploded lek or 
more dispersed population, enjoy a high 
survivorship (Frith & Frith, unpubl. data). 
Evidence fi'om sexually dimorphic polygynous 
bowerbirds, and other species, suggests that the 
strong mating skew^ in favour of older individuals 
has forced males into a long-tenn mating strategy 
involving much-delayed morphological and 
phvsiolotiical development (Beehler & Foster, 
1988; Collis & Borgia, 1992; Frith & Beehler, 
1998). 

Bower site ownership by Golden Bowerbirds is 
highly stable over years, with few successfiil 
attempts by newcomer (predominantly younger) 
males to establish themselves within bower-owning 
male society. Given this scenario, and that ex- 
perience/age has been found to play a highly 
significant role in relative male bowerbird mating 
success (Loffredo & Borgia, 1986; Collis & 
Borgia, 1992; Borgia, 1995), there is a high 
expectancy of the latter within local male Golden 
Bowerbird populations. This remains to be tested. 

ACKNOWLEDGEMENTS 

The first three years of these studies were 
peribiTned by CBF as a post-graduate student of 
Monash University, Melbourne. For this 
opportunity he is most gratefiil to Alan Lill for 
support, encouragement, advice and friendship 
and to the then Zoology Department, Monash 
University for practical help. The initial three 
years were financed in part by National 
Geographic Society Grants 1709 and 1870 to 
Alan Lill, Monash University, Andree Grilfin 
provided valued company and assistance in the 
field in various ways. We also thank Gay Crawley 
and Stephen Garnett for field assistance during 
September and October 1982. 

LITERATURE CITED 

ANDERSSON, S. 1991. Bowers on the savanna: 
display courts and mate choice in a lekking 
widowbird. Behavioural EcoIog>' 2: 2 10-21 8. 

BEEHLER, B.M. & FOSTER, M.S.' 1988. Hotshots, 
hotspots, and female preference in the 
organization of lek mating systems. American 
Naturalist 131:203-219. 

BORGIA, G 1 985a. Bowers as markers of male quality. 
Test of ah>'ix)thcsis. Animal Behav iour 35: 266-27 1 . 



340 



MEMOIRS OF THE QUEENSLAND MUSEUM 



1985b. Bower decoration and sexual competition in 
the Satin Bowcrhird (Ptilonorhynchns violaceiis). 
Behaviour, ecology and sociobiology 1 8: 9 1 - 1 00. 

1985c. Bower quality, number of decorations and 
mating success of male Satin Bowerbirds 
(Ptilonorhynchus violaceus), an experimental 
analysis. Animal Behaviour 33: 226-271. 

1986. Sexual selection in bowerbirds. Scientific 
American 254: 70-79. 

1 995. Why do bowerbirds build bowers? American 
Scientist 83: 542-547. 

BORGIA, G & GORE, M.A. 1986. Feather stealing in 
the satin bowerbird {Pi Honor hynchiis violaceus): 
male competition and the quality of display. 
Animal Behaviour 34: 727-738. 

BORGIA, G, PRUETT-JONES, S. & PRUETT-JONES, 
M . 1 985. The evolution of bower-building and the 
assessment of male quality. Zeitschrift fur 
Tierpsychologie 67: 225-236. 

BOURKE, P.A. & AUSTIN, A.F. 1947. The Atherton 
Tablelands and its avifauna. Emu 47: 87-1 16. 

BRADLEY, J.M. 1987. Vocal behaviour and annual 
cycle of the Western Bowerbird Chlamydera 
guttata. Australian Bird Watcher 12: 83-90'. 

CHAFFER, N. 1958. Additional observations of the 
Golden Bower-bird. Emu 58: 133-137. 
1984. In Quest of Bowerbirds. (Rigby: Adelaide). 

CHISIIOLM, A.H. & CHAFFER, N. 1956. Observ- 
ations on the Golden Bower-bird. Emu 56: 1-38. 

COATES. B.J. 1990. Birds of Papua New Guinea 
including the Bismarck Archipelago and 
Bougainville. Vol. 2. (Dove Publications: Brisbane). 

COLLIS, K. & BORGIA, G. 1992. Age-related effects 
of testosterone, plumage, and experience on 
aggression and social dominance in juvenile male 
Satin Bowerbirds (Ptilofiorfnmchiis violaceiis). 
Auk 109:422-434. 

COOPER, W.T & FORSHAW, J.M. 1 977. The Birds of 
Paradise and Bovver Birds. (Collins: Sydney). 

DIAMOND, J.M. 1986a. Biology of birds of paradise 
and bowerbirds. Annual Review of Ecology and 
Systematics 17: 17-37. 
1986b. Animal art, variation in bowser decorating 
style among male bowerbirds Amblyornis 
ifwrnatiis. Proceedings of the National Academv 
ofScience 83: 3042-3046. 

1 987. Bower building and decoration by the bower- 
bird Amblyoniis monmtus. Ethology 74; 1 77-204. 

1988. Experimental study of bower decoration by 
the bowerbird Amblyornis inornatus, using 
coloured poker chips. American Naturalist 131: 
631-653. 

DICZBALIS, S. 1968. Observations on the Crested 
Bowerbird Amb ly ornis macgregoriae . 
Miscellaneous Report of the Yamashina Institute 
5: 199-201. 

DONAGHEY, R.H. 1 98 1 . The ecology and evolution of 
bowerbird mating systems. (Unpubl. PhD thesis, 
Monash University: Melbourne). 



1996. Bowerbirds. Pp. 138-187. In Strahan, R. (ed.) 

Finches, Bowerbirds & other Passerines of 

Australia. (Angus & Robertson: Sydney). 
FRITH, C.B. 1989. A construction worker in the 

rainforest. Birds International I: 29-39. 
FRITH, C.B. & BEEHLER, B.M. 1998. The Birds of 

Paradise: Paradisaeidae. (Oxford University 

Press: Oxford). 
FRITH, C.B. & FRITH, D.W. 1985. Seasonality of 

insect abundance in an .Vustralian upland tropical 

rainforest. Australian Journal of Fcolo^v 10: 

31-42. 

i990a. Notes on the nesting biology of the Great 
Bowerbird Chlamvdera mwhalis (Ptilonorhyn- 
chidae). Australian Bird Watcher 13: 137-148. 

1990b. The nesting biology of the Spotted 
Bowerbird Chlamvdera macidata (Ptilonorhvn- 
chidae). Australian Bird Watcher 13: 218-225. 

i990c. Archbold's Bowerbird Archboldia papu- 
ensis ( Ptilonorhynchidae) uses plumes from 
King of Saxony Bird of Paradise Pieridophora 
alberti (Paradisaeidae) as bower decoration. 
Emu 90: 136-137. 

1993. Courtship display of the Tooth-billed 
Bowerbird Scenopoeetes dentirosfris and its 
behavioural and systematic significance. Emu 
93: 129-136. 

1 994. Courts and seasonal activities at them by male 
Tooth-bi 1 led Bowerbi rds Scennpoeete.s 
dentirosfris (Ptilonorhynchidae), Memoirs ofthe 
Queensland Museum 37: 121-145. 

1995. Court site constancy, dispersion, male 
survival and court ownership in the male 
Tooth-billed Bowerbird, Scenopoeetes 
Jt^m/>nsY/7"v (Ptilonorhynchidae). Emu 95: 84-98. 

1998. Nesting biology of the Golden Bowerbird 
Prionodura newtoniana endemic to Australian 
upland tropical rainforest. Emu 98: 245-268. 
2000a. The bower system and structures of the 
Golden Bowerbird, Prionodura newtoniana 
(Ptilonorhynchidae) on the Paluma Range, north 
Queensland. Memoirs ofthe Queensland Mus- 
eum 45(2): (this issue). 
2000b. Home range and associated sociobiology 
and ecology of male Golden Bowerbirds Priono- 
dura newtoniana (Ptilonorhynchidae). Memoirs 
ofthe Queensland Museum 45(2): (this issue). 

FRITH. C.B. & McGUlRE, M. 1996. Visual evidence 
of vocal avian mimici^ by male Tooth-billed 
Bowerbi rds Scenopoeetes dentirostris 
(Ptilonorhvnchidae). Emu 96: 12-16. 

FRITH, C.B.. BORGIA, G & FRITH, D.W. 1996. 
Courts and courtship behaviour of Archbold's 
Bowerbird Archboldia papuensis in Papua New 
Guinea. \h\s 136: 153-160. 

FRITH, D.W. 1984. Foraging ecology of birds in an 
upland tropical rainforest in north Queensland. 
Australian Wildlife Research 1 1 : 325-347. 

FRITH, D.W. & FRITH, C.B. 1 990. Seasonality of litter 
invertebrate populations in an Australian upland 
tropical rainforest. Biotropica 22: 181-191. 



BOWER ATTENDANCE AND BEHAVIOUR BY GOLDEN BOWERBIRDS 



341 



GILLIARD, E.T. 1969. Birds of Paradise and Bower 
Birds. (Weidenfeld & Nicolson: London). 

GREENWAY, J.C. Jr 1935. Birds from the coastal range 
between the Markham and the Waria Rivers, 
northeastern New Guinea. Proceedings of the 
New England Zoological Club 14: 15-106. 

HUNTER, CP. & DWYER, P.D. 1997. The value of 
objects to Satin Bowerbirds Ptilonorhynchus 
violaceus. Emu 97: 200-206. 

HYLAND, B.M.P. & WHIFFIN, T 1993. Austt-ahan 
Tropical Rain Forest Trees - an interactive ident- 
ification system. Vol. 2. (CSIRO: Melbourne). 

LENZ,N.H.G 1993. Behavioural and reproductive biology 
of the Regent Bowerbird Sericulus chrysocephaliis 
(Lewin, 1808). (Unpubl. PhD thesis, Griffith 
University: Brisbane). 
1994. Mating behaviour and sexual competition in 
the Regent Bowerbird Sericulus chysocephaJus. 
Emu 94: 263-272. 

LILL, A. 1974a. Social organization and space utiliz- 
ation in the lek-forming White-bearded Manakin, 
M. manacus Mnitalis Hartert. Zeitschrift fur 
Tierpsychologie 36: 513-530. 
1 974b. Sexual behaviour in the lek-forming White- 
bearded Manakin (M manacus trinitalis 
Hartert). Zeitschrift fiir Tieipsychologie 36: 1 -36. 

LOFFREDO, C.A. & BORGIA, G 1986. Male court- 
ship vocalisations as cues for mate choice in the 
Satin Bowerbird {Ptilonorhynchus violaceus). 
Auk 103: 189-195. 

McDONALD, D.B. 1989a. Correlates of male mating 
success in a lekking bird with male-male 
cooperation. Animal Behaviour 37: 1007-1022. 
1989b. Cooperation under sexual selection: Age- 
graded changes in a lekking bird. American 
Naturalist 134: 709-730. 

MACKAY, RD. 1989. The bower of the Fire-maned 
Bowerbird Sericulus bakeri. Australian Bird 
Watcher 13: 62-64. 

MACKAY. R.D. & CHEESEMAN, G 1 990. Extra-bower 
display of Macgregor's Bowerbird Amblyornis 
macgregoriae. Muruk 4: 63-64. 

MARSHALL, A.J. 1950. The function of vocal 
mimicry in birds. Emu 50: 5-16. 
1954. Bower-birds their displays and breeding 
cycles - a preliminary statement. (Oxford 
University Press: Oxford). 



PECKOVER, W.S. 1970. The Fawn-breasted Bovver- 
bird (Chlarnydera cerviniventris). Proceedings 1969, 
Papua New Guinea Scientific Society 21 ; 23-35. 

PRUETT-JONES, M.A. & PRUETT-JONES, S.G. 
1982. Spacing and distribution in Macgregor's 
Bowerbird (Amblyornis macgregoriae). 
Behaviour, Ecology and Sociobiology 11: 25-32. 

PRUETT-JONES, S.G. & PRUETT-JONES, M.A. 
1985. Food caching in the tropical frugivore, 
Macgregor's Bowerbird {Amblyornis 
macgregoriae). Auk 102: 334-341. 
1994. Sexual competition and courtship disrupt- 
ions: why do male bowerbirds destroy each 
other's bowers? Animal Behaviour 47: 607-620. 

ROBINSON, F.N. & CURTIS, H.S. 1996. The vocal 
displays of the lyrebirds (Menwidae). Emu 96: 
258-75. 

SCHODDE, R. & TIDEMANN, S.C. 1988. Reader's 

Digest complete book of Australian birds. 

(Reader's Digest: Sydney). 
SNOW, D.W. 1976. The web of adaptation: bird studies 

in American tropics. (Collins: London). 
1982. The Cotingas. (British Museum (Natural 

History): London). 
STILES, F.G. 1992. Effects of a severe drought on the 

population biology of a tropical hummingbird. 

Ecology 73: 1375-1390. 
THORPE, W.H. 1985. Mimicry, vocal. Pp. In Camp- 
bell, B. & Lack, E. (eds) A dictionary of birds. 

(Poyser: Calton). 
TRACEY, J.G 1982. The vegetation of the humid 

tropical region of North Queensland. (CSIRO: 

Melbourne). 

VELLENGA, R.E. 1980. Distribution of bowers of the 
Satin Bowerbird at Leura, N.S.W., with notes on 
parental care, development and independance of 
the young. Emu 80:97-102. 

VESELOVSK Y, Z. 1 978. On the biolog>' and behaviour 
of the Great Grey Bowerbird, Chlamydera 
nuchalis. Journal fiar Omithologie 119: 74-119. 

WARHAM, J. 1962. Field notes on Australian bower- 
birds and cat-birds. Emu 62: 1-30. 

ZAHAVI, A. & ZAHAVl, A. 1997. The handicap 
principle, a missing piece of Damin's puzzle. 
(Oxford University Press: Oxford). 



HOME RANGE AND ASSOCIATED S0CI0B10L0( = \ ^, i [i )LOGN or MALI- 
GOLDEN BClfWERBIRDS PRfONODURA NEfpVmNA U^TlLONOftHYNCHIDAli) 



CLIFFORD B. PRITM AKt> 0AWN fRUTl^ 

Krilh. L .B. &. \ ni\\ D.W. 200{) 06 30: I Ionic- lange imd associatcdsociohiology .irt*i ecology 
of njalc Ooltl^ Bow^rbirds Prionodura newitmiaaa (Ptilonorhynchidac). Mumain v/^q 

Tfome runge and social iateractlons of male Oolden BO\verbircih. l^nonuditr^ }li^t(^afi&. 
during display seasoais and other months of the ywr, prior to and during hower ownership, 
were studied tor six years (1 978-1 984) in upland rainforesi. Display season latis August 
to Duceraber, the wpt seaaoa rafes tarrtottng activity at fa6ff6t$. A brief pfltpiod of 
posi-mt>uIt activity otCliWfed hi lafff "M^ch to eai I v Ma> . A iradttioBal bOtVCI OWnCTlMSVer 
occupied more than one traditional bovver site simultaneously or consseciltlVely. Ttuditional 
bower sites were usually allended b> traditional (>2 seasons) bowiS" owriers. In a few 
inst.^nces (n = 3 ) the di'^^ippearnncc of a long-term traditional site ow«cr resulted ixi it being 
tcirtporaiiK visited toi a seaborn, si h> immature males, until one hecnmc established as its 
now owiitT, Similari\. n rudimcniarx bowci ;-.!ic (n 4) wns cslvihlished near (50-150ai) a 
liadjtiiHial site ifihe latter lost its long-term (uvncr The riidimciiUuy siic was (hen irreyulartj- 
allended by immature males, until one became the new owner ofthc adjacent traditional site, 
Vouny bowerless males actively attended bowt-r site--; during peak displax scnson (3S" <i of 
sightings of them) and during the post-peak disjitay seiLsini of Maicli lo eavtv Miiv (45"ii of 
sightings). From five to two years before attaining Irjdiiional bower ownciship, immature 
males visited many bower STtes,-aianavetagedi^.tunee of.i'-) I m fmin the thi;> evcninally 
came to occupy. The year before attaining full bower site occupanc) i-hi>, distance 
diiniiiished, to an average 186tn, as older inimature-plumaged birds focused then activitiBd 
nearer the siteihpy woidii eventualh OCOMW*. Most males were aduU-i»lumaged byih? ^SCOfi 
they came to QU^tiv^y dodnpy a.i3»|a[jl]Cttia) sit&fuU time,' bat afovvivel^fn Mim^tUit^b-aduft 
plumage. 

Titiidltionfil owners lelt bowers to Jbrage, bathe, drink, and collect slicks anddlBCorQtions. 
Th^ foraged)Vlailv6lv«)Qse(mean= 110m)Totheithdvt^^awi^pp^ 
noigl^oiU'ineii^¥itiW9:0n occasion. They visited o%eirben9Qra .(m^Q^^ I91m), 
ttioslly (ft2*Vy during the display season, lo steal decorations. The njjSitti^«tance travelled 
fiTOin bower -sites for all purposes averaged 143m, With a median ^!2lm (n = 152). 
Flslimaicd mean home lange size of eight" aduh mules was 7ha (range 3-lOha). □ Goyen 
Bow^rbir(i, Priomclura ncwroniana, Piilomrhynchidav. home nmges, bower acqwsttion, 
SQclsiimemctlanS' 

Clifford Bt F}ith h'ritit Honorary Research Felloyvs of ihc Queensland 

Museum, ^Prhmoduni POBnx Malwida 4SS5. AustraUa: H Sleptembcrl'J'^'j 



,lnieTe:$l in |?Qwerb|xds (Ftil{>nprh^nel;id3ifi) has 
aceisuaJecerrtfevlvtift diteto ftfeirSigttificatiBetts 
JtWcBes of the evolution of social systems and 
s©«talselection theoiy (Donagheyj. 1981; l^ruett- 
Joned*?hSett-Jones. 1982; J983; Borgia, 19^; 
frilh &FHth 1993. 1095;LeBfie 1993), Bower 
Slructures and building behaviour of male 
'bdV^ erbirds provide opportunities to examine the 
cVDlutit^ pf symbolicj e^ciBrnaliscd, Siwcondary 
S^ai cliaracters (Frith & Pnth, IW3; 1^99?!: 
iJprgia, 1995), the evolution of culturally trans- 
mitted trails (Diamond, J.V^Oii) and origins of 
aie^etfc <senser in animsils (I^^eMond. 1982, 
19S6b). In view of almost no knowledge of its 
biology and the potential significance to 
th^oreticsi Mg^f^mtioxts of atiijtidl social 



oargjuu^ations an J mating, systems m general, and 
toTh^ sA'CfliWioti of -ssBtie in bowerbirds par- 

licuiiirly. wc began studying Golden Bawcthirds 

Prkvioihiru new i(.'/n(inu, ill 1978. 

We examined (^O iradiiional bower siit% 
involving a tola I ol ^>'S> main (dccitruted) bower 
structures, during 1 978- 1 9?7 oji tlie PalumaRangc, 
aiid -^f^s^nted tt eohtpArative and qtianiified 
re\ ie\\ of them (Frith & Fritli, 2000a). A trad- 
itional bower was a large, well-established, 
structure buili at a traditional .site that was 
regularly attended, rumnlaipcd and decorated 
throughout ^bsie^etit 5edS(Bi5 by its traditiohat 
adult male owiicr. We also described short-tcnn. 
mditnentary, bowers built by inOTaiiirc inaies at 
bower sites estabfiabed lusEtr lo tradidoaal (Mie&^ 



344 





i860 m- 



+ 30 



Ridge 



120 



• 2 



• 4 



•5 



• 15 



Road 



#27 



+ 44 



+ 045 

+ ,42 (10m) 



21 (15m) 



#29 (38m) 



FTG. 1. Dispersion of 15 (including 3 just outside) traditional (•) and 10 (including one on the perimeter) 
rudimentar>' (+) male Golden Bowerbird bower sites within 50ha (1 x 0.5km) study area 1 (SAl). Only 
rudimentary sites at which marked individual birds were sighted are numbered. Faint lines indicate 
seasonal drainage gullies. Grid marks are indicated at each 200m within the 1km southern boundry 
on the figure. Bower sites just on/outside the perimeter of S A 1 are plotted within the boundary line 
and their actual distances outside it indicated. 



(Frith & Frith, 2000a), a scenario observed in 
other bowerbirds (Vellenga, 1970, 1980; 
Donaghey, 1981; Pruett- Jones & Pniett- Jones, 
1982; Chaffer, 1984; Borgia, 1986; Lenz, 1993; 
Frith & Frith, 1994, 1995, 1999a; Frith et al, 
1996). 

During the display season (late August- 
December), bower-owning males perch above 
their bowers and give loud advertisement song 
and other vocalisations including fme vocal avian 
mimicry, maintain and decorate their bowers, 
evict rival males and display to potential mates 
(Frith & Frith, 2000b). Advertisement song 
consists of a prolonged, pulsating rattle. Other 
calls include squeals, screeches, wolf-whistles, 
scolds, ft'og- and cicada-like notes, given as 
single notes, or as a medley together with fme 
vocal avian mimicry of >22 model species. Males 
leave their bower site to harvest sticks and 
decorations, and steal decorations from rival 
males for their bower, and to bathe and forage 
(Frith & Frith, 2000b). They are predominantly 
frugivorous but also eat beetles, cicadas, and 
spiders (Frith, 1989; Donaghey, 1996; Frith & 
Frith, unpubl. data). Availability of fixiits, in both 
time and space, may affect profoundly the home 
ranges of bowerbirds, but data detailing home 
ranges of fixed-point courting promiscuous male 



frugivorous bowerbirds are available only for the 
Satin, Ptilonorhynclms violaceus (Donaghey, 1981 ), 
Macgregor's (Pruett-Jones & Pruett-Jones, 
1983) and Tooth-billed, Scenopoeetes denti- 
rostris (Frith et al., 1994) bowerbirds. 

The present study provides the first inform- 
ation on distances travelled by traditional 
bower-owning male Golden Bowerbirds away 
from their bower sites. It also describes for the 
first time activities of immature males prior to 
becoming the owners of traditional bowers. 
Seasonal activities at, and away from, bowers 
during different times of the year are discussed. 
Data on male attendance levels and their 
time-budgeted activities at bowers are presented 
elsewhere (Frith & Frith, 2000b). Results of 
long-term studies of male survival, histories of 
bower ownership by individual males, and 
acquisition of adult plumage will appear 
elsewhere. 

METHODS 

STUDY AREA. This study was carried out in 
upland rainforest, at about 850m asl, on the 
Paluma Range, 7km from Paluma Township and 
80km north of Townsville, northeastern 
Queensland. The study area (19°00^S, 146°10'E) 
wasaSOhplot, 1 x 0.5km, permanently gridded 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



345 



with metal stakes (Fig. 1). For a detailed 
description of this area, see Frilh & Frith (2000a). 
The present study deals mostly with the 
ownership and movements of male Golden 
Bowerbirds based at 15 traditional and 4 
rudimentary bower sites within, or just outside 
the perimeter of our study area (Fig. 1 ). hi a few 
instances, however, birds marked as immature 
males and resighted within our study area 
eventually took up ownership of traditional 
bower sites (n = 6) beyond it. For the location of 
these bower sites (numbers 16. 22, 23, 24, 33 & 
34) see Frith & Frith (2000a: fig. 2). 

DEFFNITIONS. Definitions of bower sites and 
structures appear in Frith & Frith (2000a), and of 
the display season and vocalisations in Frith & 
Frith (2000b). We refer to a display season by the 
year in which it started (S78, S79 etc). An 
immature male was one in female-plumage and a 
sub-adult male one with some to almost complete 
adult male plumage intruding into female 
plumage. We use 'regularly attended' to imply 
full-time seasonal attendance by traditional 
owners at traditional bowers, and irregularly 
attended' to imply part-time seasonal attendance 
by immature males at traditional or rudimentary 
bowers. All statements refer to males unless 
stated otherwise. 

HOME RANGES. Movements of individually 
marked males were examined over six display 
seasons (S78-S84) and additional months of the 
year. Intensive fieldwork was from 1 August 

1 978 to 28 February 1 98 1 , save 1 May to 1 8 July 

1979 when we were absent (but relative bird 
activity at bowers was then assessed by Andree 
Griffin). In studying Golden and other bowerbird 
species during this period, 1 547 hours were spent 
carrying out systematic work: 788h during four 
hour random walks, 572h during fixed transect 
walks, and 1 87h of random searching for nests 
(see Frith, 1984; Frith & Frith 1994, 1995, 1998). 
Of the total 1547h, 893h were during display 
season months August-December; 282h during 
.lanuar^^-February when it was excessively wet 
and/or males were moulting (Frith & Frith, 
unpubl. data); 268h during March-May when 
there was some post-moult activity at bowers; and 
104h during winter months June-July. We also 
spent 343h mist-netting at bowers, as well as 
marking/retrapping individuals elsewhere during 
a standardised avifaunal netting programme 
(Frith & Frith, unpubl. data). 

Fieldwork continued through the display 
season months of S81-S84, and non-display 
season months of June 1981, February, May and 



August 1982, June 1983 and March 1984. During 
S82 and S83 we made obser\'ations at six bower 
sites in September-November 1982 for 369h, and 
November-December 1983 for 102h. Sightings 
of marked birds at bowers are presented here, but 
levels of attendance at bowers and time-budgeted 
activities at them appear elsewhere (Frith & 
Frith, 2000b). We continued netting at bower 
sites and for a general avifaunal netting pro- 
gramme during S81-S84. 

Each captured bird was metal banded and with 
a unique two colour band combination ( = 
marked). Biometrical, morphological and moult 
data will appear elsewhere (Frith & Frilh, impubl. 
data). Band colours on bower-owning birds were 
confirmed each season by direct observation at 
bowers, to avoid disturbance by retrapping, 
although many birds were retrapped opportun- 
istically away from bowers. Band colours were 
also noted during bird sightings at and away from 
bowers during other fieldwork. The present study 
deals with 26 continued males, first marked 
during S78-S8 1 in adult (n = 1 1 ) or immature (n = 
15) plumage. Numbers of sightings of marked 
individuals at bower sites were totalled for the 
months of August-December, January -February, 
April-May and May-July (S78-S84), in view^ of 
the above. 

RESULTS 

BO\VER SITES AND EXISTING OCCUPANCY. 
Traditional Bower Sites with Traditional Owners. 
Fifteen traditional bower sites were monitored 
during S78-S84: 12 within the study area and 3 
(7, 2 1 and 29) just beyond it (Fig. 1 ). All save site 
21, which was abandoned after 5 seasons, 
remained active tliroughout this study. Twelve of 
the 15 traditional bower sites were regularly 
attended over different seasons by a total of 20 
individual traditional bower owners. An aduU 
never occupied more than one traditional bower 
site, simultaneously or consecutively. 

Traditional Bower Sites Lacking Traditional 
Owners. Three of the 15 traditional bower sites 
(5, 15 and 21) lacked traditional owners 
throughout the study (Fig. 1 ). The bower at site 5 
was a large and traditional one but those at shes 
15 and 21 were rudinientar}' structures. These 
three bowers sites were irregularly attended by 
immatures, as follows: 

During S78-S84 we made 114 sightings of 
female-plumaged individuals (43 sightings of 19 
marked males, and 71 of unmarked birds whose 
behaviour indicated they were male) at sites 5, 1 5 
and 2 1 . Most (62%) of the 1 14 sightings were at 



346 



MEMOIRS OF THE QUEENSLAND MUSEUM 



traditional bower site 5, with fewer sightings at 
the rudimentary bowers of traditional sites 15 
(14%) and 21 (24%). These data are biased, 
however, because we visited site 5 most often, 
because more consistent activity was to be ob- 
served there. During these sightings we observed 
immature birds giving single call notes (25 
times), continuous medley calls with mimicry 
(33 times), and advertisement song topical of 
traditional bower-owners, only 9 times. The latter 
song was only heard during the display season, 
but other calls, although mostly (57%) heard 
during August-December, were also siven during 
JanuaV-February (19%o), March-April (19%) 
and May-July (5%). 

At these traditional sites, young males re- 
arranged bower sticks and/or decorations, or 
brought in new ones obtained elsewhere. Bowers 
had few (<1 0) decorations, in part because neigh- 
bouring adult traditional bower-owners stole 
them (confirmed by five observations of such 
thefts). Up to four young males often visited such 
sites together, where they called, displacement- 
chased each other whilst flicking/fluttering their 
wings, and sometimes perfonned brief display 
postures. 

Rudimentary Bower Sites Lacking Traditional 
Owners. Four rudimentary bower sites were 
established near to (mean = 1 04, range 30- 1 50m) 
traditional bower sites during S78-S84 and were 
active for one (site 42), two (site 25) or three 
non-consecutive (sites 30 and 44) seasons. 
Another rudimentary site (28) was 150m from 
traditional site 31 (both sites being outside SAl 
beyond traditional site 6 — see Figs 4D & 6A). 
Tliese rudimentar}' sites were irregularly attend- 
ed by immatures, but adults visited them 
occasionally, presumably to steal decorations. 

Rudimentary bower sites were usually lirst 
established near a traditional site after the 
disappearance of the latter's owner. For example, 
the adult owner of traditional site 31 had been 
badly injured (base of his rear skiall bald, torn and 
bleeding) prior to being seen at his bower on 29 
November 1978. The next day we netted and 
marked him at nearby rudimentary site 28, where 
we had sighted a sub-adult and two immatures on 
several previous occasions. We saw this adult 
male again in April 1979, his head now healed, 
being chased about his own traditional site by an 
unmarked immature. We did not see the healed 
adult again, and in S79 an unmarked adult 
occupied traditional site 3 1 and the adjacent rudi- 
mentary site was abandoned. 



In October 1979 we marked an apparently 
diseased adult at aidimentary site 42, who may 
have been the present/previous owner of nearby 
traditional site 45 (Fig. 1 ). This bird's plumage 
was in poor condition and he had a swollen 
(21.5mm long x 14.6mm diameter) hard fatty 
'bag', possibly an infected preen gland, above his 
central tail feather bases. He was not seen again, 
and an immature (black/red) took over traditional 
site 45 during the latter part of S79. 

Rudimentary bowers were usually established 
during the non-display months of March-July, by 
immatures which had previously limited their 
activities to the area surrounding the adjacent 
traditional site. Sometimes two or three im- 
matures temporarily attended such a rudimentary 
bower site. By the beginning of the following 
display season ( August'Seplcmber) the immature(s) 
would challenge the owner of the adjacent trad- 
itional site for occupancy, and by late October/ 
early November the victor exclusively occupied 
it. The rudimentary site was then abandoned. 

During S78-S84 we made 30 sightings of 
female-plumaged individuals (15 subsequently 
confirmed males and 15 unmarked birds that 
were probably male) at four rudimentary bowers 
(sites 25, 30, 42 and 44; see Fig. 1 ). During these 
sightings we heard medley calls w ith mimicry 14 
times, and advertisement song only 3 times. We 
saw adult males at rudimentary sites 5 times, but 
did not confirm if they visited them to steal 
decoration. 

ACQUISITION OF A TRADITIONAL BOWER 
SITE. Fifteen males were marked as immatures; 
1 3 were tlrst captured at an average of 3 1 7 ± 
243m (range ll0-900m) from, and two at, the 
traditional bower site they subsequently came to 
occupy. Of these 15 individuals, 14 were first 
captured within the study area: 8 at/near site 5; 2 
at site 21; 1 at rudimentary site 44 (sites lacking 
traditional owners); and 3 at sites 1,3 and 6 (sites 
with traditional owners). The initial capture sites 
of each of these 14 individuals are indicated in 
Fig. 2. The 1 5th bird (pink/red) was first captui'ed 
at a tradhional site outside the study area, but 
subsequentlv visited bowers within it (see Fia. 
3D). 

Males in Adult Plumage. Nine of the 1 5 marked 
immatures had acquired adult-plumage before 
first occupying a traditional bower site. The year 
in which two of these nine became adult was 
unknown (orange/orange and green/white; see 
Fig. 2). Six of them were in their first year of adult 
plumage when acquiring a traditional bower site. 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



347 




FIG. 2. Schematic plan to scale of localities at which 14 immature male Golden Bowerbirds were captured and 
marked within study area SA 1 . The traditional site they subsequently came to occupy and the distance to it from 
the point of their first capture are indicated- Note: b = blue; g ^ green; k ^ black; n ^ pink; o = orange; p = purple; r 
= red; w = white; * = capture sites not at bower sites; • ^ traditional bower sites with traditional owners; x = 
traditional bower sites lacking traditional owners; + = rudimentary bower site. Bower sites actually 
located immediately beyond tlie SAl perimeter are here plotted within the boundary line (see Fig. 1 
for their distance beyond it). 



and another had been adult plumaged for one 
year, as follows: 

Black/white was marked at site 5 in March 
1980 as an immature. During March-July 1980 
he visited traditional sites with traditional 
owners, including site 27 (which he later came to 
own), as well as sites 5 and 1 5 lacking traditional 
owners (Fig. 3A ). During S80 he was seen at/near 
site 5 and also at site 6, and its nearby rudi- 
mentary bower site 30, while a challenge was 
underway for ownership of traditional site 6 (see 
below). Over the next tliree years (1981-83) we 
saw him four times, the last being at rudimentary 
site 30 when he was possibly again challenging 
for site 6. In August 1984 he was the new owner 
of traditional site 27, 110m from site 5 where 
marked 4.5 years previously (Fig. 3A). 

White/blue was marked at site 5 in April 79 as 
an immature. During April 1979- July 80 he 
visited sites with traditional owners, including 
site 29 (which he later came to own), as well as 
sites 5, 1 5 and 2 1 lacking traditional owners (Fig. 
3B). From August 1980-December 81 he was 
mostly seen at/near site 5. Early in September 
1982, in his first year of aduk plumage, he 
regularly attended site 4 and we assumed he was 
its new owner, but later that month he was dis- 
placed by the owner of the previous season. By 



September 1 983 white/blue was the new owner 
of traditional site 29, 235m from site 5 where 
marked 4.5 years previously (Fig. 3B). 

Wliite/orange and black/purple were marked at 
site 5 in March 1980, as immatures. They sub- 
sequently took ownership of traditional sites 23 
and 33, being 580 and 670m distant from site 5, 
respectively (Fig. 2). From March 1980 to Nov- 
ember 1982 white/orange visited various bowers 
in the study area but in S84, 4.5 years after 
marking, he became the new adult-plumaged 
owner of site 33 (Fig. 3C). Black/purple was 
re-sighted twice after marking, once in March 
1980 at site 27, and again there in January 1981 
when in sub-adult plumage and in company of 
immature white/orange (Fig. 3C). In S82, 2.5 
years after marking, black/purple was the adult- 
plumaged owner of site 23. Another immature, 
pink/red, was marked in December 1978 at 
traditional site 24, beyond SAl but near sites 23 
(220m distant) and 33 (165m distant). We saw 
him six times in SAl during November 1979 to 
May 1 982, including once with immature white/ 
orange at sites 1 5 and 27 (Fig. 3D). In S83 he was 
the new adult owner of site 24, almost five years 
after being marked there. Thus, the above three 
males, often sighted together, as immatures, oc- 
cupied adjacent traditional sites when adults. 



348 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Feb 1981 • 17 



Black/white 



iAug-Dec 1980(11) 




19 



Sept 1982 20 



May 1982 X 21 



• B 










+ 




White/blue 


• 


• 




• 




A 


Aug 1980 - Dec 1981 (12) 


Apr 1979- July 1980 (19) 
































X 2 


m 










\ 










Sept1S82 (3) 






_X 15 


/ #27 






2 


21 X 


V 


• 29 S83 



•C 


+ 

■ 


3 • 


• 


/" 


^"^•^^ White/orange 

Mar1980-Nov 1982(11) 


/ 

/ 

/ 

/ 
/ 

2 • ' 

X 


2 ^ Y. ^ ^^-^ 33 (670 m) 
4 * \ \\ "^^^^ S84 

\ _2_____A ^-^^ 23 (580 m) 
15 X 27 --^ 382 

Black/purple 

J Mar 19B0- Jan 1981 (3) 



D 


+ 

• 


/I 

/' i 


• 


Pink/red 

Nov 1979 - May 82 (6) 


/ 

/ 

\ 








/ 


\ 




5x / 
• / 


\ 




15 x^—"'"'''^ • 
27 


24 (400 m) 
^"^^^^^ S83 




• 





E 




Orange/black 








Oct 78 t"" 

6 


16 (275 m) 


• 






• 






Orange/green 








Apr 79 - July 80 (44) 






• 


/l4 \ 








/ 








15 X 27 

S80 






• 





FIG. 3. Schematic plans to scale showing movements of seven immature male Golden Bowerbirds prior to them 
attaining aduh plumage and then acquiring a traditional bower site, during S78-S84 within parts of study area 
SAl . Number of sightings, months they were made and the season (S) an individual male acquired a traditional 
bower site are noted within Fig. 3A-E. Bower sites actually located immediately beyond the perimeter are here 
plotted within the boundary line (see Fig. I for their distance beyond it). Distances travelled to bower sites 
beyond the SAl boundry are indicated. • = traditional bower sites with traditional owner; x = traditional 
bower site lacking traditional owner; and + = rudimentary bower site; * = sightings made other than 
at bowers. 



Two young males took over traditional sites the traditional site 6 and was next seen in S79, as the 
year after we marked them: orange/black was newly adult-plumaged owner of traditional site 
marked in sub-adult plumage in October 1978 at 16, 275m from site 6 (see Fig. 3E). Orange/green 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



349 



was marked at site 5 in April 1979, as an im- 
mature. During March- July 1980, having just 
moulted into adult plumage, he challenged the 
traditional owner at site 27, being seen there 23 
times; including 9 in the traditional owner's 
presence (Fig. 3E). During August-October of 
S80 both adults apparently attended site 27. 
Twice in late October we saw the challenger 
chasing the owner from the immediate bower 
area, and in early November he had become the 
new owner. 

Males in Immature Plumage. Six of the 15 marked 
immatures remained in immature plumage when 
first occupying a traditional bower site, as follows: 
Blue/orange was first marked 230m from 
bower site 5 in March 1979. From then until July 
1980 we sighted him 17 times (see Fig. 4A). 
During S80 and S8 1 blue/orange remained closer 
to bower site 3, which he subsequently came to 
occupy. By May 1982 blue/orange had built a 
new bower based upon a pre-existing ten*eslrial 
subsidiary structure at site 3, 20m from the old 
one. He attained adult-plumage in 1983. 

Pink/purple, first marked in December 1981 
140m from site 3, was seen visiting site 3 and 
nearby traditionally-owned bowers several times 
(Fig. 4B). In early September 1982 he regularly 
attended bower site I before being displaced later 
that month by the owner of the previous season, 
hi S84, still in immature plumage, he was the new- 
owner of site 17. He acquired adult plumage the 
following year. 

During early S78 the bower at traditional site 6 
was attended by an adult male, that we marked 
there on 1 1 October, but was unseen again after 
1 7 November. Another male, blue/purple, orig- 
inally marked as an immature 205m from site 6 in 
Januar}- 1 979, was seen four times at site 6 during 
February-March, and once at site 5 in April (Fig. 
4C). During the winter months of 1979 a new 
mdimentary site (site 30) was established, 35m 
from site 6 (Figs 1 & 4C), and an immature 
male(s ) was attending it but we could not confinn 
if he was banded. By October immature male 
blue/purple was the new occupant of site 6, and 
no more activity was seen at rudimentary site 30 
that season. Our last sighting of blue/purple was 
in May 1979. 

Red/purple, first marked in April 1979, started 
attending rudimentary site 30 together with 
immature black/white in July 1980, when a 
challenge was again underway for owTiership of 
traditional site 6 (see above). By late November 
of S80 red/purple was the established new 



occupant of site 6 (Fig. 4D). Red/purple may have 
acquired traditional site 6 early, due to the 
disappearance of immature-plumaged blue/purple 
(see above), because he did not become adult- 
plumaged until 1984, four years after becoming 
the traditional owner of that site. The bower at 
rudimentary site 30 was not attended until June 
1983, when we saw red/purple (still the owner of 
site 6) there with black/white. Both birds were 
now in sub-adult plumage and black/white was 
possibly again challenging for site 6, but red/ 
purple remained its traditional owner. Rudimentary 
site 30 was not used again. 

Two other immatures had just acquired 
traditional sites (1 & 45) when first marked. Each 
site had a derelict bower, which had been 
abandoned for at least one season. Attendance by 
these two young males during their first nine 
months of occupancy was sporadic; as follows: 

In March 1979 we found a handftil of sticks, 
20m from the old bower at site 1 and we marked 
its immature builder purple/green. In S79, he 
added little to the new structure, but was seen at 
rudimentary site 25 (first located by us in S78) 
and at sites 7, 1 5 and 27, some 580, 625 and 770m 
from site 1, respectively (Fig. 5). By S 80 he was 
in adult-plumage and the new traditional owner 
of site L Rudimentary site 25 was abandoned. 

In December 1 979 we found a small new bower 
under construction at site 45, 30m from the 
derelict one. Its builder was immature black/red, 
who we marked at traditional site 21 (lacking a 
traditional owner) in October 1979 (Fig. 5). In 
March 1980 black/red was calling at newly 
established rudimentary bower site 42, 100m 
from site 45. By S80 he was the new traditional 
owner at site 45. Rudimentary site 42 was 
abandoned. Black/red was sub-adult in S81 and 
adult plum aged in S82. 

HOME RANGES. Of Non-traditional Bower 
Owners. Of 15 immatures, 13 did not own a 
traditional site when marked and two (purple/ 
green, black/red) were just acquiring one. Three 
of the 13 bowerless immatures (orange/black, 
green/blue and green/white) were not sighted 
after initial captnre until adult bower-owners (see 
Fig. 2). During S78-S84 we made 182 sightings 
of the remaining 10 individuals: 113 at bower 
sites and 69 elsewhere (usually foraging; Table 
1). Of the 113 sightings at bower sites: 38% were 
during August-December (courtship season), 5% 
January-Februar>' (wet season), 45% March-May 
(post-moult activity) and 12% June- July (Table I ). 



350 



MEMOIRS Of THE QUEENSLAND MUSEUM 



A * ^ 

r ^ Qd 1980- Nov last (7) 


K +30 




/ • 


k \ 


/ 


A. 3882 


/ • 

■ 




♦ 




Mar 1979 -July 1980 (17) 


■w. 


(x 5 

\ 




~ \ 


Blue/orange 


■^.27 

X 


X 


• 



c 






+ 30 




i» 6S79 






ft 


• 




BIU6/purple 




tan ~ Ant i Q7Q t7\ 








5 




m 








• 







\ 


r 


Dee 1061 -NavigBafO) 




• 


w 

V 

\^ 

'v 


• 2 


• 4 


Pink/purple 




X 





D 



30 



Red/purple 

Apcll79^JUiyflO{14) 




5 X 



V . 

♦ 27 



jtl^ ( polderi B(>\\ crbird's prior to 
umajie. during: S78-SjS4 within parts of 



FKj, 4, A-D, Schematic plans to scale showing movements of four imnus 
them occupying a traditional bower site while still wearing immature pU 

study area SAL Number of sightings, months they were made, and the season (S) an individual male acquireda 
traditional bower site are notdd. Bower sites actually located immediately beyond the perimeter are here plotted 
within the boundary line (see Fig. 1 for their distance beyond it). Distances travellecl to other bower sites b^ond 
the SAl boundary are indicated. • = traditional site witb traditional own^f ; % = tcadltional bowisr site 
lackingtradsttoiial ownd*; Md4-=tudt^^ *^sigbliiigsinade oilier flian atbowers. 



Of 113 sightings at bower sites: 58% were at 
traditional sites owned by traditional owners; 
39% at sites 5, 15 and 21, lacking traditional 
ovWiers; arid 3% at tuditheritaiy bower sites. 
These relative proportions varied, however, with 
individual bird age. Five to two years before the 
young males owned a traditional site tiiese 
propottioiis were 51, 45 and 4%, but one yeatr 
before such ownership they wefe 67, 3l and 2%, 
respectively (see Table 1). These differences 



retlecl the fact that in the year before occupying a 
traditional site, birds visited it more often, and 
particularly during March-July (24 of 25 
sightings). Between five' to two years before 
establishing bower tenure, imniatures visited 
traditional bower sites more distant (mean = 
391i!n) from their site of fiitUre occupancy than 
they did during the year immediately prior to 
occupying it (mearL= 186m). But sightings of 
birds away from bowers involved distances 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



351 



TABLE 1 . Histories of sightings of 1 male Golden Bowerbirds marked as immatures (female-plumage) during 
different periods of months from August 1 978-December 1984 at various bower site types and elsewhere. * = 
only 4 individuals were marked 5 seasons before coming to own a traditional bower site; the other 6 individuals 
were marked 3 (n ^ 3), 2 (n =^ 2) and 1 (n = 1) season before bower occupancy; ** = numbers in parenthesis are 
sightings of the young males at the bower site of their future occupancy; * * * = number of sightings at traditional 
bower sites 5, 1 5 and 2 1 without a traditional owner; = ! sighting represented up to 4 conspecifics visiting a 
bower simultaneously; see Results. 



No. of seasons 
individuals were 
marked prior to 
occupying a tra- 
ditional bower 
(no. of individ- 
uals *) 


Sightings at bower sites 


Sightings elsewhere 


At traditional bower sites 


At rudi- 
mentary 
bower sites 


At all 
bower sites 


In company 

of of a 
conspecific 


Mean 
distance 
(m) from 

bower site 
of future 

occupancy 


In total 


In company 
of a con- 
specific(s) 
**** 


Mean 
distance 
(m) from 

bower site 
of future 

occupancy 


with 
atraditional 
owner * * 


lacking a 
traditional 
owner *** 


August-December 


5 (n = 4) 


2(1) 








2 


1 


165 











4 (n = 4) 


4(2) 


5 


1 


10 


3 


408 


7 


1 


88 


3 (n-7) 





4 





4 





230 


5 


5 


315 


2 (n-9) 


10(1) 


1 





11 


2 


516 


1 





264 


1 (n - 10) 


5 


11 





16 


2 


211 


10 


2 


185 


Total/mean 


21(4) 


21 


1 


43 


8 


332 


23 


8 


201 


January-February 


5 (n = 4) 





























4 (n = 4) 


2 








2 


2 


625 











3 (n = 7) 


1 








1 





925 











2 (n-9) 


1 








1 





400 











1 (n=10) 


2(2) 








2 








1 





205 


Total/mean 


6(2) 








6 


2 


650 


1 





205 


March-May 


5 (n = 4) 


4(1) 


7 





11 


4 


356 


6 


1 


150 


4 {n = 4) 


3 


3 





5 


3 


444 


7 


2 


325 


3 {n-7) 


2 


3 





6 


1 


358 


8 





166 


2 (n-9) 


2 


6 


1 


9 


7 


250 


5 





202 


1 (n=10) 


16(15) 


4 





20 


4 


197 


4 





54 


Total/mean 


27 (16) 


23 


1 


51 


19 


316 


30 


3 


144 


June- July 


5 (n = 4) 




















4 


1 


124 


4 {n = 4) 


1 








1 


1 


220 


5 


1 


312 


3 (n = 7) 





























2 (n-9) 


1 





1 


2 





255 











1 (n=10) 


9(9) 





1 


11 





40 


6 


2 


39 


Total/mean 


11(9) 





2 


13 


1 


193 


15 


4 


165 


August-July 


5 (n = 4) 


6(2) 


7 





13 


5 


344 


10 


2 


142 


4 (n = 4) 


10(2) 


8 


1 


19 


9 


430 


19 


4 


225 


3 (n = 7) 


3 


7 


1 


10 


1 


430 


13 


5 


265 


2 (n = 9) 


14(1) 


7 


2 


23 


9 


415 


5 


2 


207 


1 {n = 10) 


32 (26) 


15 


1 


48 


6 


186 


19 


4 


123 


Total/Mean 


65 (31) 


44 


4 


113 


30 


345 


69 


15 


177 



352 



MEMOIRS OF THE QUEENSLAND MUSEUM 







+ 




Purple/green 

Mar 1 979 - Mar 1 980 


2 ^3 

• 


• 


7 










* Black/red 


• 






Oct 1979 -Mar 1980 








45 




X 15 




42 +-^^ 


^ X 21 


• 





FIG. 5. Schematic plan to scale of the movements of two immature male Golden Bowerbirds that had just 
occupied a traditional bower site while still wearing immature plumage, during S78-S84 within parts of study 
area S A 1 . Number of sightings are indicated. Bower sites actually located immediately beyond the perimeter are 
here plotted within the boundary line (see Fig. 1 for their distance beyond if). • =^ traditional site with 
traditional owner; x = traditional bower site lacking traditional owner; and + = rudimentary bower site. 



closer to the site of their tiiture occupancy, par- 
ticularly during the season prior to bower 
ownership (see Table 1). 

Of a total of 1 82 sightings, at bower sites and 
elsewhere, a marked immature was seen with 1 (n 
= 29), 2 (n = 8), 3 (n = 7) or 4 (n = 1), marked or 
unmarked, female-plumaged conspeciflcs (see 
Table 1). Twice we saw an immature together 
with an adult, other than the bower owner, at a 
bower site. Most sightings (67%) of marked im- 
matures at bower sites 5,15 and 2 1 involved two 
or more individuals, particularly during March- 
May (63% of sightings). 

Of Traditional Bower Owners. During S78-S84 
we made 152 sightings of 18 of the total 20 
individually marked traditional bower-owners of 
12 traditional bower sites in SAl. Sixty-eight 
were of birds at bower sites neighbouring their 
own, and 84 elsewhere (Table 2). Of the former 
68 sightings: 71% were at traditional sites oc- 
cupied by a traditional owner; 20% at traditional 
sites (5, 15 and 21) irregularly attended by im- 
matures; and 9% at rudimentary bower sites 
attended by young males (Table 2). It is note- 
worthy that all visits to mdimentary sites were by 
immatures in their first year of traditional 
bower-ownership (see Males in immature 
plumage). 



Most visitations (82% of 68 sightings) to other 
bower sites, particularly traditional ones with 
traditional owners (92%), occurred during dis- 
play months of late August-December (Table 2). 
We confirmed that >57% of such visits were to 
steal decorations (Table 2). Nearly all thefts were 
from adjacent sites, although a few males 
travelled further afield to steal (Fig. 6A). Thefts 
only occurred during the display season. 

Distances travelled to other bower sites and 
elsewhere varied little during different months of 
the year (summarised in Table 2). Visits by males 
to bower sites other than their own involved an 
average distance of 191m (n = 68, range 
40-488m) with a median of 195m, and to else- 
where (usually to forage) averaged 110m (n = 84, 
range 13-300m) with a median of 88m (Table 2). 
Thus, males remained closer to their traditional 
sites when foraging (or harvesting a decoration) 
than in visiting other bowers. Foraging home 
ranges rarely overlapped those of neighbouring 
male bower owners (Fig. 6B). Mean distance 
travelled from bower sites to all localities av- 
eraged 144m (n = 152) with a median of 121m. 
By plotting ail sightings of marked owners at 
eight traditional bower sites (i.e. Fig. 6A and B 
combined), and drawing polygons based upon 
the outermost for each (Fig. 6C), we estimated 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



353 



TABLE 2. Histories of sightings of 1 8 traditional bower-owning male Golden Bowerbirds at 12 traditional sites 
during different periods of months from August 1978-December 1984 at bower sites other than their own and 
elsewhere. * ^ number of sightings at traditional bower sites 5, 15 and 21 lacking a traditional owner; ** = 
decoration thett confirmed, but some other sightings probably also involved theft; *** ^ one sighting was with 
one conspecific; see Results. 





Sightings at bower sites 


Sightings elsewhere 


At traditional bovver 
sites 


At rudi- 
mentar>' 
bower sites 


At all 
bower sites 


For 
decoration 
theft ** 


In com- 
pany of a 
conspecific 
♦ #* 


Mean dis- 
tance (m) 

travelled lo 
other 

bower sites 


Away 
from their 
bower site 


In com- 
pany of a 
conspecific 
*♦* 


Mean dis- 
tance (m) 

travelled to 
other 

bower sites 


Months 


with a tra- 
ditional 
owner 


lacking a 
traditional 
owner * 


August- 
December 


45 


9 


2 


56 


39 


1 


205 


35 


4 


125 


January- 
February 


2 








2 








168 


9 





106 


March- 
May 


1 


3 


3 


7 





1 


156 


28 


1 


110 


June-July 





2 


1 


3 





2 


175 


12 





102 


Total/ 
mean 


48 


14 


6 


68 


39 


4 


191 


84 


5 


110 



mean overall home ranee to be 7ha (range 
3-lOha). 

On nine occasions we saw an adult in the com- 
pany of an immature, four whilst an adult visited 
another traditional site, and five times while 
foraging in the same fruiting tree (Table 2). While 
bower owning males vigorously displaced, and 
then chased, rival males out of their bower site, 
we recorded only one instance of apparent 
territoriality beyond bower sites. Adult males, 
from two adjacent bower sites, were aggressively 
challenging each other along a 200m length of 
narrow trail in rainforest equidistant between 
their bower sites. The birds gave scold and other 
calls, and flicked their wings in agitated fashion, 
while flying back and forth along their respective 
sides of, but not across, the trail. On another 
occasion tuo adult males foraged in the same 
fruiting tree without aggression. 

DISCUSSION 

Most traditional Golden Bowerbird bower sites 
are occupied by successive generations of adult 
males (Frith & Frith, 2000a; unpubl. data). 
Takeovers of traditional sites usually involved an 
intruding male being in his first year of adult 
plumage. In a few instances, a traditional bower- 
owner w as not replaced immediately by another 
adult male and, then, one of three things 
happened, a) A short-term rudimentary bower 
site was established close to the vacated 
traditional site, by immature males building a 
rudimentary structure there. This was abandoned 
once one male began regular attendance at the 



traditional site, b) A vacated traditional bower 
site was temporarily and irregularly attended by 
immature males who maintained the existing 
traditional bower, or built a rudimentary one. 
Such a new rudimentary bower might sub- 
sequently become a larger traditional one, as a 
new owTier regularly attended the site (Frith & 
Frith, 2000a, unpubl. data), c) But rarely, a trad- 
itional site was simply abandoned. Abandoned 
traditional sites may, however, be re-established 
during subsequent seasons. 

An immature male Golden Bowerbird wanders 
among the male population for at least five to six 
years before attaining first signs of adult plumage 
(Frith & Frith, unpubl. data). This wandering pe- 
riod possibly permits the novice to experience the 
social/sexual environment, while his female ap- 
pearance avoids stimulating aggressive responses 
from adult males. This provides a gradual ^appren- 
ticeship' into the intensely competitive male 
mating hierarchy, as has also been postulated for 
males of the sexually dimorphic polygynous 
manakin (Pipridae) and bird of paradise (Para- 
disaeidae) species (Lill, 1974a,b; Snow, 1976; 
Frith & Beehler, 1998). It has been suggested that 
the long retention of female plumage by males of 
such species might be part of a mating strategy, 
involving sexually active males thus concealing 
their reproductive status (Rohwer et al. 1980, 
Laska el al., 1992). Immature male Goldens 
moved extensively about the habitat visiting 
bower sites, mostly during display months of late 
August-December, and during the brief period of 
renewed activity that occurred in late March- 




FIG. 6. Schematic plans to scale of the movements of 18 traditional male owners of 12 traditional Golden 
Bovvcrbird bowers: A. to neighbouring bower sites; or B. to elsewhere: usually wiien foraging within part of 
study area SAI, during S78-S84. C, Approximated home ranges, produced by encompassing all bower visits 
and sightings elsewhere into polygons. Note: a number in parenthesis after a bower site number indicates the 
number of consecutive individual traditional owners during this study. Smaller typeface numbers on arrow lines 
indicate the number of movements/visits. Bower sites actually located immediately beyond the perimeter are 
here plotted within the bQundary line (see Fig^ 1 for their distance beyond it). Distances travelled to other bower 
sites beyond SAI are indicated. • « traditional site with traditional ownen x = traditional bower site lacking 
traditional owner; and + = rudimentary bower site. 



HOME RANGIES OF UMJi GOtUl;N UOWLRBIRDS 



355 



early Mky. Having cdmpldedtlietrni6ult, young 

males were also notgibhr active -M if aditional sites 
diinng Murch-ciirly May, w.hcn traditu>nal own- 
er'- wlTc Il'ns oiU-ii in LilicndaiiLf ;ii)d ctjmpiclinjj 
Ihcir mouli (Frilb ^ Fnth,^unpul>i. dat^J. One 
ycat he fore be(^^ng1U^turi«'bo^v^ <>wncrs 
liusiaKclvcs. immatures spent more time Ett tradi* 
trohal sixes occupied by traditional owtlers, 
cspct-ivilly nt the site they were challenging for 
and subsequently occupicLl Such visits inereased 
notably ditring March-JuK ol the season ofsub- 
sequcnttetiure^ tlic challenger liav in^ usually just 
9ttaine<iadaU plumage. 

Bower ftttcndancf by tradilional owners al Iheir 
cnfm ^tcs-was mainly during display and breed- 
ing months (Frith & Frith, 1998,2000b). During 

(Ills lime of year ihey frequently visited bowers of 
rival inales lo steal decorations (Table 1). (n 
recent yeiiTS bower deeoralion iheti has been 
studied intensively in otl^cr species, it ha$i- 
bieen deti^i^strated that mm mil ^^dm- 

inantlv from immediate neighbours (Roruia, 
1985:'l*ruou-Jones& Pruett-Jones. 1904; Inib^ 
\-:\xK 1904. 1095., :OOUb). Oui ol'serv- 

alions show Oolden Bowerbird:* do likewise, as 
ckmionstratcd in Figure 6. For a discus&ion on 
bower decorations wid th«r theft Eritli & 
Frtlh (2000b). 

Bowcrbirds travelled from their own bower to 
oilier bower sites to steal decorations yveraged 
191m (median I')5ml. iJistanccs covered to 
fbrage and iiurvest dceorulions^ were sliorivr 
(mean = H{)ra, mediBD = 88m). The overall 

sries Vf&s 121m. This fajlter figure fbr male 

bower-owning MacgregOffs'^ftSowerbird was S8rn 
(Prueil-Joncs'.*;: i'ruett-TOnes, 19831 and for 
C'l.ti-tnMiitiu Tnoih-billed Bovvctbirds wns s9m 
(f'ritli ct^ al, 1994). despite the latter species 
tr*vdUng longer distances to batheftJmik at 
creeks (not observed in Cioldcn Bnw erbirds by 
us) Similarly, die median disianee df KKm 
travelled by (_Mjldens to forage is more them 
Aiuble tliat (V.u\) observed in Tootlvbills (Frith 
feid., 19*M). t hese dilTerences may refloti (he 
sparsely and evenly dispersed malcs/bawer& of 
Maegregort un^ OciIden*» m contmi whb the 
densely cluinpcd. dispersion of Tooth-bills' 
(Courts. Ii euuld also indicate the relativc 
abtmoanec of of tfa&kttBr specto^ 

Male Satin Rovverbirds weic louiui lo mostly 
(82%^ ioroge vviUiin 50m of their bovs^rs during 
me breedifig: season of OGli)ber*Df<:^bl$r 



'(tdonaghcy; 1981). These ^ 1arx|^<^ in^sects 

(his period (Iian at oth<itim4-Wl'-50% of 

then Kuv embcr-Deccmber diefl. Diiriiig winter, 
how c\ ei. S I ' i, of lorajLiirm males v\crc up (o 2i\\hv 
from their liovvers. and the luithcst 350ni. Male 
Saiins do noi lonii kks, bul disptttHe boNwCrss 
linearly along forest edges at a mean inter-bower 
distiitice of Tl2m (Dortaghey 1^81) to 5(M)m 
(Vlarchanl, 1902). The laller authur wrote of 
males llnis iiaviii>: 'a tetrifoty ttfabDiit ?.0 ha'. 
Vellenga (19S0) wrote of each adult male Satin 
hpldingajerriU^ that included !he rudim$:nt^ 
bowier^fif (doTtrKnated) ytnmgermatc^. AKhnugh 
letritonaliiy beyond tlie bowc^site would be Ics:-; 
sinjjiisni^ in iliis ntoie insectivorous buvvcihiru 
than in hiijhly iVvigivorons ones (hieeliler & 
PrueU-JoucSv 1 9ii3; see below) tius requires study 
and clarifieaiinii.. 

W'c L->aii»ialed ihe meati yciU-nMiiu! luime raiijr^e 
ol (one or several consecutive) Golden Bower- 
bird male owncis of eiglit traditional bower ftiies 
to be 7htt.MpamhamciHngiejQriburra^ 
Adtilt male Tooth-bilTed BoW^rbfrdfe was 
determined to be 9.5ha. although males foraced 
over a suiallet atea (Hiilh el al., 199'^). DaU 
available lor Iixed-poinl'displayui^ poly^vnous 
and fm^^ivorous sjiecies of other passe line ^ oups 
so studied include the neotropical Manakms, lUid 
Cotingas (Coiin^idae), (Snow, 1970; Snoxv, 
1962a,b, 1992: Lill, I974a.b, 1976: McDonald, 
1989; Tlierv. I9'ni, Ptiuri v\ al.-, l')96). These 
studies iound that adult males defend a focal 
display site, or territory, while foraging ov^fitar 
more extensive nnddendt^d axea, qr bpmc vsoffi, 
Beehlcrand Pructt-Jbniefr(l^3lr!Witfwedapa^ 
dispersi'in ofaduh maie*^ in niiif pol v'j\ (unis bi'il 
of paradise species, in vs lncli tii ales are knovvn or 
presumed to be promiscuoiis .iiul [r, coiirlal fi\eJ 
poinl display sites (I-nlh Sl Ikehler, 199S). They 
related dispersion lo diets, and found that males 
of species with a predominantly arthropod diet 
were lerriloriid whilst those of predivminantly 
ii ugivorous ones were not. Obligate inseclivorcs 
dercruled e\i:lusive leniioncs, iiighly rmuiviir- 
ous species formed Icks, and species with 
inlenuediate diets d^o^Ved julcnno^iaTe paucras 
of dispersion. Thus 'Wmte -medes-^roll ^epSei 
defend their display sites, only thoSe^1l» more 
inseciivorons species defend a fotiElgttlgtWilOiy 
while males of iviui t- iVugi; orouS- pnCS- foMgC 
over an undefended home range. 

Adult male Golden Bowerbirds vigorously 
defended their bower site auainsi ri\ als, but we 
s«V too little ofaOult males louedier away trom 
bov^ to assess the nacore of mteiactiOn^ pits^ 



356 



MEMOIRS OF THE QUEENSLAND MUSEUM 



While male Macgregor's Bowerbirds aggressively 
defend bower sites, few aggressive interactions 
occurred away from Ihem (Pruett-Jones & 
Pmetl-Jones, 1982). Our fmdings suggest the 
observation that 'Although males do not defend 
territories in the usual sense, they do occupy areas 
over which they exert dominance. We do not 
mean that males exclude rivals from the area 
(beyond the bower site) or prevent them from 
foraging there, but that they do prev^ent rivals 
from establishing courtship sites' (Beehler & 
Foster, 1988) is applicable to male Golden 
Bowerbirds. Further field work is required to 
clarify the question of (undefended) foraging home 
range versus the extent of (defended) territory in 
this species. 

Fruit in tropical rainforest is largely econom- 
ically undefendable by passerine birds, as a resuh 
of its spatial and temporal unpredictability 
(Snow, 1976; Lill, 1976; Beehler, 1983; Beehler 
& Pruett-Jones, 1983; Frith & Beehler, 1998). 
Thus the loss of extensive territoriality in pre- 
dominantly frugivorous species, such as the 
Golden Bowerbird. Fruit availability in time and 
space may therefore have profound effects upon 
home ranges of bowerbirds (Beehler & Pruett- 
Jones, 1983; Frith et al., 1994). Male bowerbirds 
must remain close to their bowers if they are to 
successfully defend their structures and decor- 
ations from rivals, and attract and mate females 
(Frith & Frith, 1993). They attempt to maximise 
time spent at their bowers, as do Tooth-billed 
Bowerbirds at their courts (Moore, 1 991 ; Frith & 
Frith, 1994). This is predictable behaviour for a 
population of promiscuous males that must 
compete for opportunities to fertilise numerous 
females at a fixed focal site during a relatively 
brief mating season (Frith & Frith, 1993, 1995). 

Snow ( 1 976) considered a predominantly frug- 
ivorous diet the main pre-condition to a 
polygynous mating system, in which rainforest 
dwelling male passerines spend most of their 
time in advertising/attending/defending a 
traditional focal courtship area and performing 
elaborate courtship. Seasonal abundance of fruit 
in the habitat is theoretically so great as to enable 
males to be emancipated from nesting duties, and 
females to nest alone and unaided. In tropical 
rainforest-dwelling and predominantly 
frugivorous Golden Bowerbirds, and in many 
other ecologically similar passerines, the fruit 
diet would appear to have greatly influenced the 
evolution of a polygynous mating system, 
associated morphology, male survival and 
longevity, dispersion, focal courtship sites with 



territoriality limited to them, and undefended 
home ranges (Snow, 1976; Frith & Beehler, 1998). 

ACKNOWLEDGEMENTS 

The first three years of these stiadies were 
performed by CBF as a post-graduate student of 
Monash University, Melbourne. For this op- 
portunit>' he is most grateful to Alan Lill for 
support, encouragement, advice and friendship 
and to the then Zoology Department, Monash 
University for practical help. The initial three 
years were financed in part by National 
Geographic Society Grants 1709 and 1870 to 
Alan Lill, Monash University. For constructive 
criticism of an earlier draft of this contribution we 
thank David Snow. 

LITERATURE CITED 

BEEHLER, B.IVl. 1983. Frugivory and polygamy in 
birds of paradise. Auk 100: 1-12. 
1989. The birds of paradise. Scientific American 
261: 116-23. 

BEEHLER, B.M. & FOSTER, M.S. 1988. Hotshots, 
hotspots, and female preference in the organiz- 
ation of lek mating systems. American Naturalist 
131:203-219. 

BEEHLER, B.M. & PRUETT-JONES. S.Q 1983. 
Display dispersion and diet of birds of paradise, a 
comparison of nine species. Behavioral Ecology 
and Sociobioiogy 13: 229-38. 

BORGIA, G. 1985. Bower decoration and sexual com- 
petition in the Satin Bowerbird {Plilonorhynchus 
viaiaceus). Behaviour, ecology and sociobiolo^ 
18: 9I'W(). 

1986. Sexual selection in bowerbirds. Scientific 
American 254: 70-79. 

1 995. Why do bowerbirds build bowers? American 
Scientist 83: 542-547. 

CHAFFER, N. 1984. In quest of bowerbirds. (Rigby: 
Adelaide). 

DIAMOND, J.M. 1982. Evolution of bowerbirds' 
bowers: animal origins of aesthetic sense. Nature 
297: 99-102. 

1986a. Biology of the birds of paradise and 

bowerbirds. Annual Review of Ecology and 

Systematics 17: 17-37. 
1986b. Animal art: variation in bower decorating 

style among male bowerbirds Amblyornis 

inornatus. Proceedings of the National Academy 

of Sciences USA 83: 3042-3046. 
DON AGHEY, R.H. 1 98 1. The ecology and evolution of 

bowerbird mating systems'. (Unpubl. PhD thesis: 

Monash Universits; Melbourne). 

1996. Bowerbirds. Pp. 138-187. In Slrahan, R. (ed.) 
Finches, Bowerbirds & Other Passerines of 
Australia (Angus & Robertson: Sydney). 

FRITH, C.B. 1989. A construction worker in the 
rainforest. Birds International 1: 29-39. 



HOME RANGES OF MALE GOLDEN BOWERBIRDS 



357 



FRim C.B. & BEEHLER, B.M. 1998. The birds of 
paradise: Paradisaeidae. (Oxford University 
Press: Oxford). 

FRITH, C.B. & FRITH, D.W. 1993. Courtship display 
of the Tooth-billed Bowerbird Scenopoeefes 
dentirostris and its behavioural and systematic 
significance. Emu 93: 129-136. 

1994. Courts and seasonal activities at them by 
male Tooth-billed Bowerbirds Scenopoeefes 
dentirostris (Ptilonorhynchidae). Memoirs of the 
Queensland Museum 37: 121-145. 

1995. Court site constancy, dispersion, male 
survival and court ownership in the male Tooth- 
billed Bowerbird, Scenopoeetes dentirostris 
(Ptilonorhynchidae). Emu 95: 84-98. 

1 998. Nesting biolog> of the Golden Bowerbird 
Prionodura newtoniana endemic to Australian 
upland tropical rainforest Emu 98: 245-268. 
2000a. The bower system and structures of the 
Golden Bowerbird, Prionodura newtoniana 
(Ptilonorhynchidae) on the Paluma Range, north 
Queensland. Memoirs of the Queensland Mus- 
eum 45(2); (this issue). 
2000b. Bower attendance and behaviour at them by 
male Golden Bowerbirds, Prionodura new- 
toniana (Ptilonorhynchidae). Memoirs of the 
Queensland Museum 45(2): (this issue). 

FRITH, C.B., BORGIA, G & FRITH, D.W. 1996. 
Courts and courtship behaviour of Archbold's 
Bowerbird Archholdia papuensis in Papua New 
Guinea. Ibis 136: 153-160. 

FRITH, C.B. & FRITH, D.W. & MOORE, G 1994. 
Home range and extra-court activity in the male 
Tooth-billed Bowerbird, Scenopoeets dentirostiis 
(Ptilonorhvnchidae). Memoirs of the Queensland 
Museum 37: 147-54. 

FRITH, D.W. 1984. Foraging ecology of birds in an 
upland tropical rainforest in north Queensland. 
Australian Wildlife Research 11: 325-347. 

LASKA, M.S., HUTCHINS, M., SHEPPARD, C, 
WORTH, W. & BRLfNFNa D. 1992. Repro- 
duction by captive unplumed maleLesser Bird of 
Paradise Paradisaea minor evidence for an 
alternative mating strateg} ? Emu 92: 108-1 1 . 

LENZ, N.H.G 1993. Behavioural and reproductive 
biology of the Regent Bowerbird Sericulus 
chrysocephaliis (Lewin, 1808). (Unpubl. PhD 
thesis, Griffith University: Brisbane). 

LILL, A. 1974a. Social organization and space 
utilization in the lek-forming White-bearded 
Manakin, M. manacus trinitalis Hartert. 
Zeitschrift fiir Tierpsychologie 36: 5 15-530. 
1974b. Sexual behaviour in the lek-forming 
White-bearded Manakin (A/ manacus trinitalis 
Hartert). Zeitschrift fiir Tierpsychologie 36: 1 -36. 



1976. Lek behaviour in the Golden-headed 

Manakin Pipra aythrocephala in Trinidad (West 
Indies). Advances in Ethology 18: 1-84. 

MARCHANT, S. 1 992. A Bird Observatoiy at Moruya, 
NSW 1975-84. Eurobodalla Natural History 
Society Occasional Publication No. 1: 1-99. 

McDonald, D.B. 1989. cooperation under sexual 
selection; Age-graded changes in a lekking bird. 
American Naturalist 134: 709-730. 

MOORE, GJ. 1991. Seed dispersal by male Tooth- 
billed Bowerbirds Scenopoeetes dentirostris 
(Ptilonorhynchidae), in north-east Queensland 
rainforest: Processes and consequences, (Unpubl. 
PhD thesis, James Cook University of North 
Queensland: Townsville). 

PRUEl T-JONES, M.A. & PRUETT-JONES, S.G 1982. 
Spacing and distribution in Macgregor's Bower- 
bird {Amblyornis macgregoriae). Behaviour, 
Ecologx- and Sociobiology 1 1 : 25-32. 
1983. The bowerbird's labor of love. Natural 
Historv 9: 49-55. 

PRUETT-JONES, S.G & PRUETT-JONES, M.A. 1994. 
Sexual competition and courtship disruptions: 
why do male bowerbirds destroy each other's 
bowers? Animal Behaviour 47: 607-620. 

PRUM, R.O., KAPLAN, J.D. & PIERSON, J.E. 1996. 
Display behaviour and natural histor>' of the 
Yellow-crowned Manakin (Heterocerus 
flavivertex: Pipridae). Condor 98: 722-735. 

ROHWER, S., FRETWELL, S.D. & NILES, D.M. 
1980. Delayed maturation in passerine plumages 
and the deceptive acquisition of resources. 
American Naturalist 115: 400-437. 

SNOW, B.K. 1970. A field study of the Bearded 
Bellbird in Trinidad. Ibis 1 12: 299-329. 

SNOW, D.W. 1962a. A field study of the Black and 
White Manakin, Manacus manacus, in Trinidad. 
Zoologica47: 65-104. 
1962b. A field study of the Golden-headed Mana- 
kin, Pipra ervthrocephala. in Trinidad, W.I. 
Zoologica 47:^183-198. 
1976. rhe Web of Adaptation: bird studies in 

American tropics. (Collins: London). 
1982. The Cotingas. (British Museum (Natural 
History): London). 

THERY, M. 1990. Ecologie et comportement des 
oiseaux Pipridae en Guyane: leks, frugivorie et 
dissemination des graines. (Unpubl. PhD thesis, 
University of Paris: Paris). 

VELLENGA, R.E. 1970. Behaviour of the male Satin 
Bowerbird at the bower. The Australian Bird 
Bander 8: 3-11. 
1 980. Distribution of bowers of the Satin Bowerbird 
at Leura, NSW. with notes on parental care, 
development and independence of young. Emu 
80: 97-102. 



SOME NEW GIVETIAN (LATE MIDDLE DEVONIAN) GASTROPODS FROM THE 
PAFFRATH AREA (BERGISCHES LAND, GERMANY) 



JIRI FRYDA 

Fryda, J. 2000 06 30: Some new Givetian (late Middle Devonian) gastropods from the 
Paffrath area (Bersisches Land, Germany). Memoirs of the Oueemland Museum 45(2): 
359-374. BrisbaneriSSN 0079-8835. 

Revision of Givetian (Middle Devonian) gastropods from the Paffrath area (Bergisches 
Land, Germanv) has identified a large number of new taxa described herein. These are 
Retispira fasselli sp. nov., Stenoloron (Pajfi-atholoron) go/dfussi subgen. et sp. nov., 
Quadricarina (Blodgettiiia) reticulata subgen. et sp. nov., Kirchneriello gen. nov., 
Eiserhardtia inepta gen. et sp. nov., Cerithioides whidhornei sp. nov., Plagiothyra 
midiispiralis sp. nov., Naticopsis (Paffrathopsis) subgen. nov., Paffrathia lotzi gen. et sp. 
nov., Cookiloxa pulchra gen. et sp. nov., Palaeozygopleura (Rhenozyga) subgen. nov. and 
Heteroloxonema gen. nov. □ Gastropods, Devonian, Givetian, Germany. 

Jiri Fryda, Czech Geological Survey, Klarov 3. 118 21 Praha L Czech Republic. 
(fiyda@cgu.cz): 14 January 2000. 



During the last fifteen years the number of 
studies on Devonian gastropods has increased 
rapidly (e.g., Blodgett & Rohr, 1989; Blodgett, 
1992, 1993; Blodgett & Johnson, 1992; Kase & 
Nishida, 1986, 1988; Homy, 1992, 1994; Homy 
& lordan, 1993; Fryda, 1992, 1998c,d, 1999b; 
Fryda & Manda, 1997; Fryda & Bandel, 1997; 
Gubanovelal., I995;Cook, 1993, 1997;Cook& 
Caniilleri, 1997; Bandel & Fryda, 1998). Impetus 
for increased interest in this relatively neglected 
group of Devonian fossils were tlie seminal studies 
of Blodgett et al. ( 1 988, 1990) who identified the 
great potential utility of Palaeozoic gastropods as 
paleobiogeographic indicators. In addition, 
research focused on the higher taxonomy of the 
Devonian gastropods, based on protoconch 
morphology, has shown that during Devonian 
time a large change-over occurred from a fauna 
of typical Early Palaeozoic cast, to one of a more 
modem aspect (see Fryda, 1999a and references 
therein). Revision of the Givetian (Middle Dev- 
onian) gastropods from Gemiany (Fryda, 1998a) 
have identified a large number of new taxa and 
revised material untouched for up to 150 years 
(Goldfuss, 1844). The results of this revision 
have only partially been published (Bandel & 
Fryda, 1998, 1999; Fryda, 1998b, 1999a) and the 
majority of them have been prepared for mono- 
graphic publication. Because new taxonomic names 
established in an unpublished study (Fryda, 
1 998a) are prepared to be used in non-taxonomic 
studies by other authors, there is a problem with 
their priority as well as the fact that some of them 
would be noinina nuda. For this reason, some 
new Givetian (Middle Devonian) gastropods 



from the Paffrath area (Bergisches Land, 
Germany) are established herein. Monographic 
treatment with detailed descriptions of all species 
belonging to newly established taxa as well as the 
evaluation of their palaeogeographical and strati- 
graphical distributions is in preparation (Fryda & 
Bandel, in prep.). All described gastropods come 
from the collection of Dr Volker Ebbighausen 
and are deposited in the Senckenberg Museum in 
Frankfurt au Main (Germany). 

SYSTEMATIC PALAEONTOLOGY 

AMPHIGASTROPODA Simroth, 1906 
BELLEROPHONTOlDEAM'Coy, 1851 
BELLEROPHONTIDAE MToy, 1851 
KNIGHTITINAE Knight, 1956 

Retispira Knight, 1945 

TYPE SPECIES. Retispira hellireticulata Knight, 1945. 

REMARKS. Retispira (Knight, 1945) is a com- 
plex genus within Knightitinae later regarded as 
a subgenus oYKnightites Moore, 1941 (Knight el 
al., 1 960). Batten ( 1 972 ) suggested that Retispira 
represents a separate genus and this was followed 
by Gordon & Yochelson ( 1 987). Retispira ranges 
from the Devonian to the Permian. 

Retispira tasselli sp. nov. 
(Fig. IF-J) 

ET\TV10L0CiY. For the Australian paleontologist Cliris 
Tassell. 

MATERIAL. 6 specimens from Hen'enstrundcn. localit>' 
9, Bergisches Land, Germany (coll. Ebbighausen). 
Holotype: figured herein as Fig. 1 F,G 



360 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG \ . A-C, Stenoloron (Paffratholoron) goldfussi sp. now. A, Holotype, oblique view x 3.5; B, Holotype, apical 
view x3.6; C, Paratype, apertural view x3.8. D, E, Qiiadricarina (Blodgettina) reticulata sp. nov. D, 
Holotype, basal view showing a reticular pattern X 4; E, Holotype, apertural view X 5. F-J, Retispira tasselli sp. 
nov. F, Holotype, lateral view showing a shell ornamentation x 4.8; Q Holotype, dorsal view X 4.8; H, Paratype 
A, dorsal view x 7; I, Paratype B, dorsal view X 4.5; J, Paratype C, apertural view x 5. 



GIVETIAN GASTROPODS FROM GERMANY 



361 



T\TE LOCALITY. Bucheler Schichten, Middle Devonian 
(Givetian). Henenstrunden, Beigisches Land, Germany. 

DIAGNOSIS. Species of Retispira with small 
shell having a broad, flat selenizone raised above 
the shell dorsum; shell narrowly and deeply 
phaneromphalous; lateral portions of whorl 
profile rounded into narrow umbilici without any 
angulation; apertural lip of mature whorl is 
slightly expanded on its lateral and lunbilical 
portions; shell ornament consisting of spiral and 
collabral lirae forming a reticulate pattern of 
longitudinally elongated rectangles. 

DESCRIPTION. Small bellerophontid shell 
bearing a broad, flat selenizone raised above shell 
dorsum. Shell narrowly and deeply phanerom- 
phalous. Whorl profile slightly concave on each 
side of selenizone and gently convex in 
dorsolateral portions. Lateral portions of whorl 
profile curve into narrow umbilici without any 
angulation. Shallow and wide sinus in anterior 
margin of the outer lip culminates in a slit gen- 
erating a raised selenizone. Selenizone bounded 
by two distinct spiral threads; width of the 
selenizone about 15% of width of shell dorsum. 
Apertural lip of the mature whorl slightly 
expanded on its lateral and umbilical portions. 
Shell ornament consists of spiral and collabral 
lirae; collabral lirae widely spaced and more 
expressed than closely spaced spiral lirae; spiral 
lirae crossing the collabral lirae form a reticulate 
pattern of longitudinally elongate rectangles. 
Distances of both spiral and collabral lirae var>^ 
considerably during the ontogeny. 

REMARKS. Retispira tasselli resembles 
Retispira leda (Hall, 1861) from the Middle 
Devonian of North America in its similar whorl 
expansion in the adult whorl. However, R. tasselli 
sp. nov. ditTers by having a narrower shell, raised 
selenizone and by the absence of ornamentation 
of its selenizone. Bellerophon elegans de 
Ferussac & Orbigny, 1840 noted by Archiac &. 
Vemeuil (1842: 354, pi. 29, fig. 2) from the 
Paffrath region is similar to Retispira tasselli and 
most probably also belongs in Retispira. It is 
distinguished from the latter by the absence of a 
raised selenizone. Retispira tasselli is also 
similar to Retispira sp. of Blodgett (1992) from 
the Eifelian of Alaska, but may be distinguished 
by its more raised selenizone. 

Rollins et al. (1971 ) discussed similarities of 
Retispira and Bucampsis which have similar 
shell shapes. Merriamites Blodgett & Johnson, 
1995 {=Merriamella Blodgett & Johnson, 1992, 



previously occuppied homonyin), based on the 
Middle Devonian species Merriamites eurekae 
from Nevada, is also similar to Retispira in having 
a reticulate ornamentation. Merriamites differs 
from Bucampsis, the latter having an ornament of 
spiral threads, in having a much broader 
longitudinal keel on the inner floor of the whorl, a 
prominent reticulate pattern and a prominent, 
rounded parietal tooth. The presence ot a keel on 
the inner floor of the whorl is a character by which 
Bucanopsis and Merriamites (both are placed in 
Carinaropsinae) differ from Retispira. Similarities 
in these genera with some Knightitinae were also 
noted by Blodgett & Johnson (1992). 

ARCHAEOGASTROPODA Thiele, 1925 
GOSSELETINIDAE Wenz, 1938 

Stenoloron Oehlert & Oehlert, 1888 
Stenoloron (Paffratholoron) subgen. nov. 

TYPE SPECIES. Stenoloron (Pqffi-atholown) goldfussi 
sp. nov. 

ETYMOLOGY. Pqffi'atholoro}% for the type locality. 

DIAGNOSIS. Subgenus oi^ Stenolorofi Oehlert & 
Oehlert, 1888 having low spired, phanerom- 
phalous shell with slowly expanding whorls. 

REMARKS. Stenoloron (Paffratholoron) 
subgen. nov. has slowly expanding whorls. The 
distance of the upper suture from the shell per- 
iphery (measured in the apical view) is about 
15% of its total shell width. This value is double 
(i.e. 30%) in the generic type species, Stenoloron 
viennayi (Oehlert & Oehlert, 1888). In addition, 
Devonian species of Stenoloron Oehlert & 
Oehlert, 1888, such as Stenoloron subaequitera 
(Chapman, 1916) from the Early Devonian of 
Australia, Stenoloron minor Blodgett & Johnson, 
1992, from the Middle Devonian of North 
America, and Stenoloron pollens Perner, 1903 
from the Early Devonian of Europe have similar 
values of whorl expansion as the type species (i.e. 
about 30%). For this reason the species described 
below is placed in a new subgenus. 

Stenoloron (Paffratholoron) goldfussi sp. nov. 
(Fig. lA-C) 

ETYMOLOGY. For tlie Gennaii paleontologist August 
Goldfuss, who studied Devonian gastropods of the Paffrath 
area. 

MATERIAL. 3 specimens from Untertlial, Pallrath ai'ea, 
Beigisches Land, Gennany (coll. Ebbighausen). Holot> pe: 
figured herein as Fig. 1 A,B. 



362 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TYPE LOCALITY. Unlerer Plattenkalk, Middle 
Devonian (Givetian), Untertlial, locality 63, Beigisches Land, 
Gemiany. 

DIAGNOSIS. As for subgenus. 

DESCRIPTION. Small, turbiniform shell with a 
wide umbilicus, width about one quarter of total 
shell width. Width of shell having about 7 whorls 
measures about 15mm. Sides of shell slightly 
convex (i.e. cyrtoconoid) due to decrease of the 
pleural angle in the last two whorls. Narrow, 
slightly convex selenizone situated high on the 
whorl bounded by a single shallow groove on the 
each side. Whorl profile above the selenizone 
rounded and nearly subhorizontal at the upper 
suture; sutures are shallow. Suture just below its 
selenizone in younger whorls; in the mature 
whorl, the distance of the lower groove bounding 
the selenizone and the lower suture is about one 
half of selenizone width. Shell surface smooth. 

The initial portion of the shell is lost in the 
holotype but it is preserved in another specimen 
tVom the same locality figured herein as Fig. IC. 
The high spired early whorls distinctly protrude 
above the upper, convex shell surface. During the 
further shell growth, the pleural angle continually 
increases, but in the last two whorls it again 
slightly decreases. For this reason the sides of the 
early shell are concave and slightly convex in the 
mature shell. 

REMARKS. Stenoloron (Paffi-atholoron) gold- 
fussi sp. nov. is the only species assigned to the 
subgenus. It is distinguished from the species of 
Slerioloron (Stenoloron) by shape of its many- 
whorled shell and characteristic slow expansion. 
The selenizone in Stenoloron (Paffratholoron) 
goldfussi sp. nov. is wider than in the generic type 
species Stenoloron (Stenoloron) viennayi and 
Stenoloron (Stenoloron) subaequitera (Chapman, 
1916), but is simHar in width to Stenoloron 
(Stenoloron) minor Blodgett & Johnson, 1992. 
Stenoloron (Paffratholoron) goldfussi represents 
the youngest occurrence of the genus, previously 
reported from the Early Devonian and Silurian. 
The only other Middle Devonian species known 
is Stenoloron (Stenoloron) minor Blodgett & John- 
son, 1992 from tlie Eifelian of central Nevada. 

EOTOMARIIDAE Wenz, 1938 

Quadricarina Blodgett & Johnson, 1992 
Quadricarina (Blodgettina) subgen. nov. 

TYPE SPECIES. Quadricarina (Blodgettim) reticulata 
sp. nov. 



ETYMOLOGY. For the American paleontologist Robert 
B. Blodgett. 

DIAGNOSIS. Similar to Quadricarina (Quadri- 
carina), with low spired shell having distinctly 
raised selenizone above the whorl surface; shell 
ornament consisting of reticulate pattern. 

REMARKS. Quadricarina Blodgett & Johnson, 
1992 is represented by four Middle Devonian 
species. Three species, including its type species 
Quadricarina glahrobasis Blodgett & Johnson, 
1992, come from the Eifelian of Nevada. 
Blodgett & Johnson (1992) also include 
Pleurotomaria lenticularis Goldfliss, 1844 from 
the Givetian of Germany. The turbiniform, 
phaneromphalous shells of Quadricarina species 
are characterised by a selenizone bordered by 
two pairs of revolving cords. A gently concave 
selenizone is situated at mid-whorl height. Shell 
oniament consists of fine collabral ribs. Quadri- 
carina (Blodgettina) subgen. nov., is similar to 
hitherto known species of Quadricarina 
(Quadricarina) in general shell shape and the 
position of the selenizone, but differs in having a 
distinctly raised selenizone above the whorl sur- 
face and reticulate shell ornament. 

Quadricarina (Blodgettina) reticulata sp. nov. 
(Fig. 1D,E) 

ETYMOLOGY. In reterence to the presence of reticulale 
ornamentation. 

TYPE LOCALITY. U. Plattenkalk lunestones. Middle 
Devonian (Givetian), Unterthal, locality 63, Bergisches 
Land, Germany. 

MATERIAL. 10 specimens from Unterthal, locality 63, 
Bergisches Land, Germany (coll. Ebbighausen). Holotype: 
figured herein as Fig, 1 D,E. 

DL^GNOSIS. As for subgenus. 

DESCRIPTION. Small, turbiniform shell with a 
selenizone situated at shell periphery. Mature 
spire consisting of about six whorls, width 
approximately i3mm. Outer lip of aperture forms 
a shallow sinus culminating in a peripheral carina 
that gives rise to a flat selenizone; selenizone 
distinctly protruding above whorl surface. 
Selenizone bordered by two pairs of revolving 
cords; outer pair forming its margin and inner 
pair spaced between outer cords. Inner cords 
always slightly thinner than the outer cords. Very 
naiTow groove between the inner cords. Upper 
and lower whorl surfaces convex; sutures deep 
and impressed. Whorl profile below selenizone 
fornis a rounded shell base with a deep, funnel- 



GIVETIAN GASTROPODS FROM GERMANY 



363 



like umbilicus. Width of umbilicus about one 
quarter of total shell width. Wliorls embrace 
below the selenizone. Distance between lower 
margin of selenizone and lower suture greater 
than width of the flat, protruding selenizone. 
Shell ornament above and below spiral and 
collabral threads forming a reticulate pattern. 
About 25 spiral threads below the selenizone 
fomi small tubercles where they are crossed by 
collabral threads. 

REMARKS. Quadricarina (Blodgelt'ma) retic- 
ulata subgen. et sp. nov. may be distinguished 
from all species of Ouadricarwa (Quadricarma) 
by the presence of peripheral carina bearing a 
selenizone and by its reticulate ornamentation. 
However, it resembles Quadricarina sp. nov. B 
of Blodgett & Johnson (1992) in its low-spired 
shell, wide umbilicus and slightly raised, carinate 
periphery but differs by having the selenizone 
distinctly situated above the suture, and having 
reticulate ornament. Wliorls embrace sliglitly above 
the selenizone in Quadricarina sp. nov. B. The 
carinate periphery suggests that the poorly known 
Quadricarina sp. nov. B may belong to Quadri- 
carina (Blodgettina) . 

RAPHISTOMATIDAE Koken, 18% 
OMOSPIRJNAE Wenz, 1938 

Kirchneriella gen. nov. 

TYPE SPECIES. Turbo striatus Hisinger in Goldlliss, 
1844. 

ETYMOLOGY. For the Gennan paleontologist Heinrich 
Sylvester Kirchner, who in 1915 published 'Mittel- 
devonische Gastropoden von Soetenich in der Eifel'. 

DIAGNOSIS. Turbiniform, relatively low-spired 
shell with a veiy indistinct selenizone situated 
high on the whorls; lateral side of whorls 
rounded; shell ornament consisting only of spiral 
cords; spiral cords on the phaneromphalous shell 
base generally regularly spaced; more closely 
spaced than those on the upper whorl surface; a 
spiral cord is also present on the selenizone 
surface. 

REMARKS. Kirchneriella gen. nov. closely 
resembles the Permian Callistadia Knight, 1945 
in general shell shape, position of the indistinct 
selenizone and shell ornament. Carboniferous 
and Permian species of Callistadia may be 
distinguished from Kirchneriella gen. nov. by the 
shape of the lateral whorl profile which is 
rounded in Kirchneriella, but nearly vertical and 
straight or gently concave in Callistadia (see 



Knight, 1945; Hoare & Sturgeon, 1978). 
Kirchneriella also shows some similarities with 
the type species of Gyroma Oehlert, 1888, 
Pleurotomaria haconnierensis (Oehlert, 1888), 
from the Lower Devonian of France, however 
this species has collabral and spiral ornaments, in 
contrast to Kirchneriella. Nevertheless, some 
Devonian species assigned to Gyroma, like the 
Frasnian Gyronui altaica (Vemeuil, 1845) and 
Gyroma subcapillaria Vostokova, 1 966, lack col- 
labral ornament and probably belong to 
Kirchneriella, Blodgett (1992) noted the first 
occurrence of pre-Carboniferous Callistadia in 
Middle Devonian rocks (Eifelian of Alaska). Un- 
fortunately, only two, poorly preserx ed specimens 
are known, preventing determination of whether 
they belong to Callistadia or to Kirchneriella. The 
Middle Devonian 'Turbo ' mult istricttus Kirchntr, 
1915 and 'Turbo ' aequistriatus Kirchner, 1915 
may belong to Kirchneriella, but need to be 
resludied in detail. Kirchneriella striata 
(Hisinger in Goldfuss, 1844) is the only known 
species of this genus. 

Family UNCERTAIN 
Eiserhardtia gen. nov. 

TYPE SPECIES. Eiserhardtia inepta sp. nov. by monotypy. 

ETYMOLOGY For the Gemian paleontologist Klaus 
Eiserhardt for his kind help with SEM photos. 

DIAGNOSIS. Discoidal shell having a very wide 
and ilat selenizone which forms the whorl 
periphery; shell base phaneromphalous; width of 
umbilicus about one quarter of the shell width; 
selenizone of younger whorls is situated in 
distinct, U-shaped groove close to upper suture; 
upper suture forms abapical margin of the seleni- 
zone; shell ornamentation consisting of spiral and 
collabral elements forming a reticular pattern. 

REMARKS. Eiserhardtia inepta sp. nov. is 
distinguished by its U-shape groove opening of 
the selenizone on the preceding whorl (Fig. 2G). 
In addition, the very wide and flat selenizone 
forming the whorl periphery is not common among 
Palaeozoic gastropods. Eiserhardtia resembles 
some Palaeozoic genera like the Ordovician 
Latitaenia Koken, 1 925, Chepultapecia Ulrlch in 
Weller & Clair, 1928, and Liospira Ulrich & 
Scofield, 1897 and the Devonian Umbotropis 
Pemer, 1 903 in general shell shape. However, the 
unusual subsutural groove easily differentiates it 
from all these genera. 



364 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2. A-C, KircJmeriella striata (Hisinger in Goldfuss, 1 844); A, apertural view x 4.5; B, basal view showing a 
spiral ornamentation, same shell as A, x 3.6; C, apical view x 5. D, Cerithioides whidbornei sp. nov., lateral 
view X 3.8. E-G, Eiserhardtia inepta sp. nov., Holotype; E, apical view showing shell ornamentation x 1 1 ; F, 
apertural view X 13; G, oblique view showing a subsutural groove x 13. 



GIVETIAN GASTROPODS FROM GERMANY 



365 



Eiserhardtia inepta sp. nov. 
(Fig. 2E-G) 

ETYMOLOGY. Latin, ineptiis^ inept unable. 

MATERIAL. Only one complete shell, Unterthal, locality 
63, Bergisches Land, Gennany (coll. Lbbighausen). 
Holotype: figured herein as Fig. 2E-G 

TYPE LOCALITY U. Plattenkalk limestones. Middle 
Devonian (Givetian), Unterthal, locality 63, Bei^ischcs 
Land, Gennany. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Medium-sized, dextrally coiled, 
discoidal shell with a wide and flat selenizone 
situated at the shell periphery. Mature spire with 
about four whorls; width more than double its 
height. Outer lip of aperture forms a shallow sinus 
culminating at a peripheral selenizone. Selenizone 
parallel with shell axis and ornamented by 
irregularly spaced lunulae. Width of flat seleni- 
zone about one quarter of the whorl height. 
Outline of aperture elliptical with longer axis 
roughly perpendicular to shell axis. Whorl profile 
above selenizone slightly convex, except close to 
the upper suture where it forms a U-shaped 
groove. Selenizone of younger whorls situated in 
this U-shaped groove (Fig. 2G); upper suture just 
below selenizone. Width of umbilicus about a 
quarter of the total shell width. Shell ornament 
consists of spiral and collabral thi eads foniiing a 
reticulate pattern. About 15 regularly spaced, 
spiral threads on the shell above the selenizone 
are crossed by backward curving collabral 
threads. 

MURCHISONIIDAE Koken, 1 896 

Cerithioides Haughton, 1 859 

TVPE SPECIES. Cerithioides telescopium Haughton, 1 859. 

REMARKS. Cerithioides telescopium, was 
described from the Carboniferous of Ireland. 
Donald (1892) placed in its synonymy two 
additional Carboniferous species, Murchisonia 
maxima and GIvptohasis cotnccu both earlier 
described by de Koninck (1883). Batten ( 1966), 
who revised the type species of Cerithioides^ 
expressed the opinion that Cerithioides coniciis 
(de Koninck) is an independent species having 
Cerithioides maximus (de Koninck) as a junior 
synonym. He also agreed with Donald's 
placement of M//r(:7?/50/?/a (Cerithioides?) oweni 
Donald, 1895 into Cerithioides and additionally 
placed Cerithioides? gleanensis Longslaff, 1926 
into synonymy with this species. Batten (1966) 



placed Murchisonia eversolensis Stauffer, 1909 
from the Middle Devonian of Ohio into 
Cerithioides and considered it to be the earliest 
representative of the genus. Small shells of 
Cerithioides incomptum Linsley, 1 968 from the 
Middle Devonian of the North America probably 
do not belong to Cerithioides. Cerithioides 
whidhornei sp. nov. has a similar whorl profile to 
C. eversolensis (Staulfer, 1 909) and also lacks the 
spiral costae on shell base. Both species may 
belong to a new subgenus within Cerithioides 
using the latter characters differing them from the 
Carboniferous Cerithioides species. 

Cerithioides whidbornei sp. nov. 
(Fig. 2D) 

ETYMOLOGY. For the British paleontologist GF Whid- 
bome who contributed much to our knowledge of the 
Devonian gastropods. 

MATERIAL. 10 specimens from the type locality (coll. 
Ebbighausen). Holot>'|">e: figured herein as Fig. 2D. 

TYPE LOCALITY. Middle Devonian (Givetian) lime- 
stones, Bergisch Gladbach, locality 65, Bergisches Land, 

Germany. 

DIAGNOSIS. Species of Cerithioides lacking 
spiral costae on the shell base; wide selenizone 
situated in the middle of whorl. 

DESCRIPTION. High-spired, dextrally coiled 
shell with almost straight sides. Whorl profile 
distinctly convex close to both upper and lower 
sutures and nearly flat or slightly concave at the 
mid-whorl. Suture moderately deep. Shell base 
rounded and anomphalous. Wide selenizone 
situated at mid-whorl with width about 20% of 
the distance between the lower and upper sutures. 
Outer apertural lip bears a wide sinus culminating 
in a wide slit. Margins of apertural sinus fonn an 
angle of about 90°. Inner lip covered by thin 
inductura. Ornamentation consists only of growth 
lines. Selenizone limited by a fine, spiral groove 
on each side and ornamented by irregularly 
spaced lunulae. 

REMARKS. Cerithioides whidbornei sp. nov. 
may be distinguished from Cerithioides ever- 
solensis by the position of the selenizone. In 
C. whidbornei, the wide selenizone is situated 
mid-whorl, however, in C. eversolensis (SXmffcr, 
1909) it is distinctly below mid-whorl. Its upper 
margin runs closely to mid-whorl and the distance 
between its lower margin and the lower suture is 
smaller than the width of the selenizone (see 
StautTer, 1909, pi. 16, fig. 13). Cerithioides 



366 



MEMOIRS OF THE QUEENSLAND MUSEUM 



whidhoniei sp. nov. may be also distinguished 
from the type species C lelescopium Haughton, 
1859, by the absence of spiral costae on its shell 
base. Cerithioides oweni Donald, 1895, differs 
from both Devonian species and the type species 
by its whorl profile which has a distinct 
angulation below the selenizone {see Donald, 
1895: pi. lOJigs 1, la). 

NERITIMORPHA Golikov & Starobogatov, 
1975 

NERITOIDEARafinesque, 1815 
PLAGIOTHYRIDAE Knight, 1956 

Plagiothyra Whidbome, 1892 

TYPE SPECIES. Monodonta purpurea Archiac & 
Vemeuil, 1842. 

REMARKS. Whidborne (1892) established 
Plagiothyra for species Monodonta purpurea 
Archiac & Verneuil, 1842 and his new species 
Plagiothyra archon Whidbome, 1 892. The latter 
was described from the Middle Devonian of 
southwest England. Vostokova ( 1 966) suggested 
a placement of the species Palaeotrochus prae- 
cursor Clarke, 1 885 into Plagiothyra, but earlier 
Vostokova (1961) had established Pseudoplagi- 
othyra for that species. Tw'o new undescribed 
species of Plagiothyra occur in the Devonian of 
the North America (R.B. Blodgett, pers. comm.); 
one in the Eifelian age Cheeneetnuk Limestone 
of the west-central Alaska and a second species in 
the Emsian age Disappointment Bay Fonnation, 
Canadian Arctic Islands. The last species is the 
first Early Devonian occurrence, and thus the 
oldest representative of Plagiothyra. 

Plagiothyra multispiralis sp. nov. 
(Fig. 3E-H) 

ETYMOLOGY, muliispiralis, referring to the shell 
ornamentation fbmied by many spiral threads. 

MATERL\L. More than 30 specimens from the Unterthal, 
locality 63, Bergisches Land, Germany (coll. 
Ebbighausen). Holotype: figured herein as Fig. 3mL 

TYPE LOCALITY. U. Plattenkalk limestones, Middle 
Devonian (Givetian), Unterthal, locality 63, Beigisches 
Land, Gemianv'. 

DIAGNOSIS. Species of Plagiothyra ornamented 
by many spiral threads; plate-like ridge bearing a 
triangular tooth within aperture smaller than in 
type species. 

DESCRIPTION. Dextrally coiled, turbiniform 
shell with a rapidly expanding whorls, shell 
height about 15mm. Shell base anomphalous. 



sutures shallow and adpressed. Whorl profile 
shows distinctly stronger convexity near upper 
suture where it fomis a small ramp. Whorls join 
the preceding whorl below its periphery. The 
margin oi' the outer apertural lip passes down- 
ward from the upper suttu'e with strong backward 
obliquity, but without any curvature. Columellar 
lip covered by a thick inductura. Prominent, 
triangular tooth occurs on a very low platelike 
ridge in the middle of the parietal wall. Shell 
ornamentation consists of many spiral threads 
that may be crossed by line growth lines. Number 
of spiral threads between the sutures is about 20. 
Threads more distinct on adapical part of outer 
whorl surface. 

REMARKS. Plagiothyra multispiralis sp. nov. 
differs from the type species, Plagiothyra 
purpurea Archiac & Verneuil, 1 842, by having a 
lower-spired shell that is ornamented by many 
spiral threads. Shell ornamentation on 
Plagiothyra purpurea consists of several distinct 
rows of tubercles. The number of spiral elements 
between the sutures is about 20 in Plagiothyra 
multispiralis sp. nov., but only 8 in Plagiothyra 
purpurea. Shell ornament on P. multispiralis 
resembles that of the other two genera in the 
family Plagiothyridae, Dirachis Whidbome, 
1891, and Littorinides Knight, 1937. However, 
shells of the Middle Devonian Dirachis, being 
ornamented by a lower number of more distinct 
spiral elements, bear two teeth in the aperture. 
Tlie Carboniferous Littorinides has one low tooth 
on the columellar lip in contrast Plagiothyra 
multispiralis sp. nov. The similar shape of the 
apertural tooth and its position within the 
aperture, as well as the similar shell shape of P. 
purpurea and P. multispiralis, necessitate 
placement ot'the new species in the Plagiothyra. 

NERITOPSIDAE Gray, 1847 
NATICOPSINAE Miller, 1889 

Naticopsis M'Coy, 1844 
Naticopsis (Paffrathopsis) subgen. nov. 

TYPE SPECIES. Natica suhcostata Archiac & Verneuil, 
1842. 

ETYMOLOGY. For the type area, Paffrath, Germany. 

DIAGNOSIS. Naticopsis having moderately low 
spired shells with strong collabral cords extend- 
ing across entire whorl surface. 

REMARKS. Yochelson (1953) placed Natica 
suhcostata Archiac & Verneuil, 1842 into 
Naticopsis (Jedria) based on the Carboniferous 



GIVETIAN GASTROPODS FROM GERMANY 



367 




FIG. 3. A-D, Naticopsis (Paffi'athopsis) subcostata (Archiac & Verneuil, 1 842); A, abapertural view of shell with 
well preserved colour pattern x 3.3.; B, same shell covered with ammoniuin chloride X 3.3; C, apical view X 3; 
D, apeitural view x3. E-H, Plagiothyra multispiralis sp. nov.; E, Paratype A, apertural view showing a 
triangular tooth on low platelike ridge of the parietal wall X 7; F, Paratype B, apertural view X 3.5. G, Holotype, 
apertural view X 5; H, Holotype, abapertural view X 5. 



368 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Naticopsis meeki Knight, 1933. Naticopsis 
(Jedria) subcostata was the only Devonian 
species placed into Naticopsis (Jedria). Blodgett 
(1992) described Naticopsis (Jedria) decker i 
from the Eifelian (Middle Devonian) of Alaska 
which is the oldest representative of this 
subgenus. Gordon & Yochelson ( 1 987 ) noted that 
the body whorl extended downward rather than 
outward, and that the swelling on the outer whorl 
face above the periphery indicates the subgenus 
Jedria. All these shell characters may be seen in 
the Carboniferous species o f Naticopsis (Jedria) 
as well as in Naticopsis (Jedria) deckeri, but not 
in Natica subcostata Archiac & Vemeuil, 1842. 
A^. subcostata has a moderately low spired shell 
with strong collabral cords extending across the 
entire whorl surface. These shell characters 
distinguish this species trom all species included 
in Naticopsis (Jedria) as well as from other sub- 
genera of Naticopsis. Naticopsis (Pqffrathopsis) 
subcostata is the only species currently known. 

Naticopsis (Paffrathopsis) subcostata 

(Archiac & Vemeuil, 1842) 
(Tig. 3A-D) 

non Buccinites suhcosiatus, Schlotheim 1 820: 130; 

Schlotheim 1822: 63, pi. 12. fig. 3. 
Natica subcostata, 1842: 366, pi. 34, figs 5, 5a, 6; Goldfliss 

1844: 116. pi. 198. fig. 22. 
Turbonitella subcostata (Archiac & Verneuil) Lotz 1900: 

212; Kirchner 1915 (partim): 238; Paeckelmann 1922: 41. 

MATERIAL. Five complete shells and several fragments 
from the Unterthal, locality 63, and one complete shell 
from Beigisch Gladbach, locality 76, Berglsches Land, 
Germany (coll. Ebbighausen). 

REMARKS. Goldfuss (1844) synonymised the 
species Buccinites subcostatus Schlotheim, 1 820 
■dxxd Natica subcostata Axchvdc 8l Vemeuil, 1842. 
However, Schlotheira's figure (see Schlotheim, 
1822, pi. 12, fig. 3) of Succinites subcostatus 
ditTers trom that of Natica subcostata (compare 
with Archiac & Vemeuil, 1 842, pi. 34, figs 5, 6). 
Material from Paffrath evidently belongs to the 
species Natica subcostata Archiac & Vemeuil, 
1842, which is the first unquestionable illustration 
of this species. For this reason, Archiac & Ver- 
neuil's name is considered to be a valid species 
name. 

Colour Pattern. Traces of original colour pattem 
in Naticopsis (Paffrathopsis) subcostata 
(Archiac & Vemeuil, 1842) were obser\^ed in 
specimens from Unterthal, locality 63, and 
Bergische Gladbach, locality 76. The geometry 
of the colour pattem in specimens from the both 
localities is the same. The colour pattern consists 



of three spiral bands of black irregular spots (Fig. 
3 A). The distance between the spots in each band 
is about equal to their diameter. The whorl 
surface close to the upper suture is without any 
colour. Archiac & Vemeuil (1842: pi. 34, figs 5, 
6) and Roemer (1876: pi. 32, fig. 8) figured the 
same colour pattem as described herein in their 
figures of Natica subcostata {=Naticopsis 
(Paffrathopsis) subcostata). 

Paffrathia gen. nov. 

TYPE SPECIES. Paffiathia lotzi sp. nov. 

ETYMOLOGY. For the type area, PaflErath, Gemiany. 

DIAGNOSIS. Small, low spired turbiniform 
shell ornamented by strong, sharp collabral ribs 
extending across the entire whorl surface; shell 
wall very thick; whorl profile suboval. 

REMARKS. Characteristic shell omament con- 
sisting of the distinct collabral ribs (Fig. 4A, B) 
distinguishes Paffrathia from all Palaeozoic 
members of the family Neritopsidae. Absence of 
spiral elements and the presence of distinct col- 
labral elements of the shell ornamentation have 
complicated the subfamily-level position of 
Paffrathia gen. nov. as well as Naticopsis 
(Paffrathopsis) subcostata (Archiac & Vemeuil, 
1842) within the Neritopsidae. According to 
Knight et al. (1960) theNaticopsinae unites shells 
without omament (except for subsutural collabral 
threads or cords in some species). On the other 
hand. Palaeozoic members of the Neritopsinae 
are omamented with pustules. Only the type 
species Paffi'athia lotzi sp. nov. is known. 

Paffrathia lotzi sp. nov. 

(Fig. 4A,B) 

Turbonitella sp.?, Lotz, 1900: 213, pi. 3, fig. 8. 

ETYMOLOGY. For fl. Lotz who originally figured this 
species. 

MATERIAL. One complete shell from the Unterthal, 
locality 63, Bergisches Land, Gemiany (coll. Ebbig- 
hausen). Holotype: figured herein as Fig. 4A,B. 

TYPE LOCALITY. U. Plattenkalk limestones, Middle 
Devonian (Givetian), Unterthal, locality 63, Bergisches 
Land, Germany. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Small, low spired turbiniform 
shells with at least four whorls. Whorl profile 
convexly arched. Height of the adult shell 
approximately equals width. Whorls embrace 
near base. Irmer whorl profile suboval, sutures 



GIVETIAN GASTROPODS FROM GERMANY 



369 



very deep. Whorl profile runs from the upper 
suture laterally for short distance to rounded 
spiral angulation, far from this angulation it 
curves downward fonning convex lateral part of 
the whorl, and then continues across the rounded 
whorl base into deep umbilicus. Outer apertural 
lip is prosocline. Shell wall in adult whorl is 
relatively thick; about 20% of inner apertural 
diameter. Shell ornament consists of strong, 
sharp collabral ribs extending across the entire 
whorl surface. Ribs regularly spaced and their 
distance roughly equals half of the diameter of 
the aperture. Fine growih lines may be obser\ ed 
on the surface between collabral ribs. 

REMARKS. Paffrathia lotzi is a rare element of 
gastropod fauna of tlie Givetian (Middle Devonian) 
U. Plattenkalk limestones of Germany, with the 
illustrated specimen (Fig. 4A,B) the only shell 
available for study. The specimen figured by Lotz 
(1900: pi. 3, fig. 8) seems to belong to this 
species. However, a larger collection of this 
species is needed for the study of shell variability. 

? LOXONEMATOIDEA 
Koken, 1 889 

Cookiloxa gen. nov. 

ETYMOLOGY. For the Australian paleontologist Alex 
Cook. 

TYPE SPECIES. Coofdloxapulchra sp. nov. 

DIAGNOSIS. Slender, regular, high-spired and 
multiwhorled shell with straight sides having a 
gently convex whorl profile; sutures deep, whorls 
more arched close to both upper and lower 
sutures; external surface of whorls ornamented 
by collabral costae which are asymmetrically 
bent and crossed in the mid-whorl by one spiral 
costa. 

REMARKS. Cookiloxa resembles Ausiraloxa 
Cook & Camilleri, 1 997 in its shell ornamentation. 
The latter genus is based on Australoxa tasseUi 
Cook & Camilleri, 1997 from the Middle Dev- 
onian of Australia. The angular whorls of A. 
tasseUi are ornamented by prominent opistho- 
cline ribs and a low, sharp sinus is developed on 
the angular whorl periphery (see Cook & Camilleri, 
1997: fig. 9E-L). One spiral cord occurs just 
above the lower suture. Cookiloxa gen. nov. 
differs from Australoxa in having a rounded 
whorl profile, a much smaller shell and by the 
absence of a peripheral sinus and lower spiral 
cord. Cookiloxa also resembles the Pemiian 
genera Loxosonia Batten, 1985 and Cibecuia 



Winters, 1963 in shell ornament. Batten (1985) 
interpreted the spiral groove-like element of their 
shell ornamentation as a selenizone. Cookiloxa 
differs from Cibecuia by a different whorl 
profile, being distinctly convex in contrast to the 
latter genus which has a flat whorl profile. 
Cookiloxa has a similar whorl profile as both 
known species of Loxosonia. i.e. Loxosonia 
horrwotoma Batten, 1985 and Loxosonia zy^o- 
pleuroides Batten, 1985. Cookiloxa may be 
distingushed from both species of Loxosonia by 
the position of its spiral element of shell orna- 
mentation. In Cookiloxa costa-like spiral element 
is situated in the middle of the whorl, but in 
Loxosonia the groove-like spiral element (?= 
selenizone) occurs close to the upper suture. 
Cookiloxa also resembles the Ordovician genera 
Spiroecus Longstaff, 1924 and Donaldiella 
Cossmann, 1903. Spiroecus LongstalT, 1924, 
differs from Cookiloxa in having a more prom- 
inent spiral ornamental element being situated 
above the mid-whorl and forming a distinct 
angulation (see holotype in KnighU 1941; pi. 48, 
fig. 10). In addition S. girvanensis differs from 
Cookiloxa pulchra by the absence of collabral 
costae (but see specimen figured by Wenz, 1938: 
fig. 929b). Cookiloxa may be distinguished fi'om 
the genus Donaldiella by its difiering shape of the 
whorl profile and outer apertural lip. The Middle 
Devonian Cookiloxa pulchra sp. nov. from 
Germany is the only known species. 

Cookiloxa pulchra sp. nov. 
(Fig. 4C,D) 

ETYMOLOGY. Latin, /Wc/?^/; beautiful. 

MATERIAL. About 30 specimens from the Herren- 
strunden, locality' 9, Bergisches Land, Gennany (coll. 
Ebbighausen). Holotype: figui'ed herein as Fig. 4D. 

TY?E LOCALITY. Bucheler Schichten, Middle Dev- 
onian (Givetian), Herrenstainden, locality 9, Betgisches 
Land, Gemiany. 

DIAGNOSIS. As for genus. 

DESCRIPTION. Small high-spired, dextrally 
coiled shell, up to 13 whorls present. Sides of 
shell nearly straight, whorl profile distinctly 
convex; maximum convexity^ close to both upper 
and lower sutures. Whorl profile nearly flat at 
mid-whorl. Sutures moderately deep. Lateral part 
of the whorl curves uniformly to base part, form- 
ing a smooth curvature without any angulation. 
Shell base rounded and anomphalous. Shell 
ornamentation consists of asymmetrically bent 
collabral costae being crossed in the mid-whorl 



370 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 4. A, B, Pqffrathia lotzi sp. nov., Holotype; A, apertural view x 4.5; B, apical view x 4.5. C, D, Cookiloxa 
piilchra sp. nov.; Paratype A, lateral view X 14; D, Holotype, lateral view x9. E, F, Palaeozygopleura 
(Rheinozyga) retrostriatum (Kirchner, 1915) comb, nov.; E, lateral view showing the regularly spaced 
orthocline costae x 1 5; F, lateral view of larger shell x 9. G-I, Heteroloxonema moniliforme (Goldfuss, 1 842) 
comb, nov.; G, apertural view x 3.4; H, lateral view x 3.4; I, oblique view showing an early whorl x 125. 



OlVE'riAN CiAiiTRUPODS FROM UURMANY 



371 



by one spiral costa. Begjnniiigattlic upper sulurea 
the coltabral q9sta&tusti iliptpsoclinc direction^ 
gradually curve Jtt mi Oftiteclfne direction, mul 
Ihen inopistlutclinc directum near the nnd-whori 
rorming a \\ndc, asymniclhcal cirt:lii.-d siriiis 

RHMARkS. Shape o\' ihc collahral costtic in 
Cookilo\''f pn/i hrn sp hdv icsciTibIt; those a[' 
some species q{ f^almo^yRopieura Horny, 1955 

/! (it^'onicam (Pcmcr, !*)()7) and the Middle 
Dc\oniaii, P hutttLlinnac (Haii, 1861)). Spiral 
coslLi HI L'^okiinxK piilchra is mosi probablv jusi 
elements ul* the sliell ornaiucniuiiou and not a 
ncUTow stieni/onc. Without knowletiJg^ of the 
initial |taitx)t'ti^ sIikcI] Cof^ldtom jgeiv^^ftv^its 

PALAEOZYOOPLEURIDAE 
HcMrn^, 1955 

rjl:K'n/v:::oplcuraHoniy, 1955 
l'ahK'ij/y;:opicur.i (Kheiio/\i»a) subgcn iiov. 

TYPE !iP£CIES. Laxont-'ma fvtfv.siha(um Kirchtia; 

lliL- n\cr Rhine. 

DIAGNOSIS. Fcjluco:ygoplenrct w iili i u ihoc i ii it; 
or slightly opisthocyit collabral costac; whorl 
between sutured Strongly Hrched; ^hell uyrto- 
-conoid or with sti'atght sides. 

REMARKS. Palaeozygophur^ (Khi^t^^^^gvf 
aubgcn. nov. difllrs liom iili ot]icrsubgC^iia.ol" 
Palaeoz}[i^op/t'Nnt hv Mu sti ipc of collabral 
costae which are orilir.chnc nr slightly opit;iho- 
cyrt. HahieozygiJpIcura ifihcfUKv^a) may be 
distinguished by its strongLy arched whorls 
wiii^art: also hi'^erJtteQ4i£iaMoJh^sybgenera 
of Pittaeozy:^opfettf*a. The orthoclinc or slightly 
Lipisthocyrt slupc of costatr rescinbic that of 
Pevoiiozyi^a Hom^, 1^55. The whorls in 
Oevotwzyga Uom^. 1^)55 are disiincily 
shouldefecland intliiS ix^ian region flattened, in 
contrast to thdl frt Vlit Fuiniozv^npleura 
(Rhemzyga) suhgcn. nov. which has strcinuly 
arched whorl sides. F^lodgetl ( 1^)92) deserihed a 
new genus Maskozygoplctira based on the 
F, i I c I i a ii A Ins A ozy^oph'tiyu crass i cosiatti 
BlodgcU. \ W2 which he tentatively placed in the 
family Pscudqjygoplcuridae Knight^ 1930. 
Pataeozygopieitra (Rhemozyga) subgen. nov. 
rescmhlcs AfiiskozygopleNtci in Its shell omainent- 
iilion, hul the lormer ta\on may be disiiagui&hed 
irom the latter its^much nairowec^idl and circular 



aperture. Blodgett(I992) noted broad vatiabilily 
i}f )a& Alaskoiygopleura a-amcostata^ but all of 
#he figured shells of bis species have GVOld, yd- 

apieall} aarR>wer apertures, considerably w ider 
shells and higher whorls than Palaeozygopk'UfV 
(Rhemzyga) retmatl^XtUm (Kirchncr, 1§I5). 

Untbrtunalely Uiere are np -dataaboitt the early 
shell in the Qivctian s>XiZz\^%t}tf Palae(>z\x<>i>ieura 
tRhfimzyijii} subuen. nOV, All oilier .^ubyenerj of 
Pc(laeoz\y,(fi'ic(tni haVC protoeoneli^. of ihc 
archaeogasiropod lype (Frydy ik Bandel, H'V 
The piacemeni of Rhenozyga in the genus- 
P^ktmzygopleum shtuild beprt^d by ty pe ofils. 
protoconch. Besides llie type species, Palaeo- 
zygtiplcum fRheno2yga) retrostriatum (Kirchrier^ 
l*)15 l. several other 'I^uv/ztw/ 'species froni the 
Midiile Devonian ot Germany like Lunnnna 
spira)jjohosiwi Kirchner, 1915 may belong \o the 
new Palu^x^gc^leuro (Rb^nps}f&^, Pala^fjzyMO 
fmtrv n^it^ayfOobK. 1^7 trom ^ )mc^ 
"Devonian of Australia also is trans forred to the 
new PiiLtL'icyooplciira /Rhcnoz)'gah A detailed 
study mI'-jII Miduic I jca onian species resembling, 
the genus PaluL<:\ yj'p/cura as well as cam- 
pati^Ott with the 1 lielian Afft^0zygr»p!etfr^' 
CfossicoyfdUt Blodiicu, 199': r-, in preparaliotv. 

? HI- i I Kt tv ! R( ii>! iA J richer, lli85 
Iteteroloxonernst gen. nov 

TYPE SPECIES. Tnrntalta monmonms Goldtuss. I.S^M. 

E IA MOLOGY. \ combination of die genene naiue 
Loxonemu and the prefix ht'ten>- because of the presumed 
heteroslrophic coiling ofils early shell. 

DIAGNOSIS. Slender, high-spired and multi- 
whorled shell; first whorl planispirally or slightly 
sinistrally coiled; shell apexbltmtand dome-like; 
teleoconch sides slightly^ conV^Jifr; whorl wofile 
gently convex between shallow, butwithmstinet 
sutures; leleconeh surtace smooth. 

Rl^MARKS. I hc smooth tcleoctmeh of //c/nvj- 
ht\nmnut gen lesendiles iliat o\ intmy 

loxonc*matids inclucUng Loxom'mo, but may b^? 
distinguished hy itsdomc-Iikc shell apex tbrmcd 
by plamspirally or slighlly sinislrally eoiletl eaHy 
whorl. All known Devonian loxonematoidetm 
ga'^lrop'.'ds sucli i\s Kaloptychia Pemer. 1*^07, 
Stylonema Perner, 1907, PalaeozygopU'tJrft 
(Paiaeo^gopreUrt^ Horny, 1955, PaliUi>- 
^jmpleuro fPala^&zyga) Horny, 1955, aAd 
Pafoeozygopl^pa (Bohemozyga) FT^aa i 
Bandel. [99? have an acute shell apet which i*i 
tormcd by the axchaeogastropod-typcnrotOGonch. 
Ji»kfro}mtmeim^.mv. h^iVtg apiattispiralty 



372 



MEMOIRS OF THE QUEENSLAND MUSEUM 



or slightly sinistrally coiling of its early whorl 

may be distinguished by this character from all 
known loxonematoideans. This character dis- 
tinguishes Hetewloxonema gen. nov. also from 
the genus Donaldina Knight, 1933, based on the 
Carboniferous Aclisina gran/onensis Donald, 
1898. According to Knight (1941), the latter 
species has ""earliest two whorls free from the 
spire, the first whorl planispirally coiled and 
tilted', in contrast to the blunt and dome-like shell 
apex of Hetewloxonema. In addition, the shells 
of the type species are ornamented by numerous 
spiral lirae (see Knight 1941: pi. 48, fig. 3a-e), 
Hetewloxonema moniliforme (Goldftiss, 1844) 
from the Middle Devonian of Germany is the 
only known species. 

Heteroloxonema moniliforme 

(Goldfriss, 1844) 
(Fig. 4G-I) 

Tumtella moniliformis, Goldfliss 1844, p. 103, pi. 196, fig. I. 
Loxonema monilifomui (Goldfuss), Bandel 1994. pi. 4. fms. 
9-10. 

Donaldina moniliforma (Goldfuss). Nutzel,1997: 209, pi. 34 

s-r. 

MATERIAL. 13 complete shells from the Unterthal, 
locality 63, Pafirath area, Gennany (coll. Ebbighauscn). 

DIAGNOSIS. As for genus. 

DESCRIPTION. Medium sized, high-spired, 
dextrally coiled shells having at least 9 whorls. 
Shell sides slightly convex; sutural slope approx- 
imately lO"". The first whorl planispirally or 
slightly sinistrally coiled. Shell apex blunt and 
dome-like. Whorl profile gently convex and 
slightly tlaltened in median region; width of 
whorl almost twice of its height. Sutures are 
shallow but distinct. Shell base anomphaious, 
teleoconch surface smooth. 

REMARKS. Bandel ( 1 994) figured the early whorl 

in Heteroloxonema moniliforme (Goldfuss, 
1844) which is, according to his observation, 
sinistrally coiled. For this reason he considered it 
to be a member of the Heterostropha. If his in- 
terpretation is correct, then Heteroloxonema 
moniliforme represents the oldest known 
member of this subclass (Bandel, 1 994) and also 
the only heterostrophid gastropod older than the 
Frasnian/Famennian mass extinction. Never- 
theless, no protoconch/teleoconch boundary was 
hitherto observ ed in this species. 

ACICNOWLEDGEMENTS 

I thank Dr Volker Ebbighausen for providing 
material from his collection. This publicafion is 



based on my studies of the Siluro-Devonian 

gastropods supervised by Klaus Bandel 
(Universitat Hamburg), partly sponsored by the 
U.S. -Czechoslovak Science and Technology 
Joint Fund in cooperation with the Czech 
Geological Survey and U.S. Geological Survey 
under Project Number 95057. I gratefiilly thank 
Robert B. Blodgett (Oregon State University) 
and Alex G. Cook (Queensland Museum) for 
their useful critical remarks to the manuscript and 
improving its language. 

LITERATURE CITED 

ARCHIAC, E.J.A.D. & VERNEUIL, E.P.D. 1842. On 
the fossils of the older deposits in the Rheinish 
Province. Transactions of the Geological Society 
of London 6: 303-410. 

BANDEL, K. 1994. Triassic Euthyneura (Gastropoda) 
from Sl.Cassian Formation (Italian Alps) with a 
discussion on the evolution of the Heterostropha. 
Freiberger Forschungshetl C 452: 79-100. 

BANDEL, K. & FRYDA, J. 1998 Position of 
Euomphalidae in the system of the Gastropoda. 
Senckenbergiana Lethaea 78 (1/2): 103-131. 
1999. Notes on the evolution and higher classifi- 
cation of the subclass Neritimorpha (Gastropoda) 
with the description of some new taxa. Geologica 
et Paiaeontologica 33: 219-235. 

BATTEN, R.L. 1966. The lower Carboniferous 
gastropod fauna from the Hotwells limestone of 
Compton Martin, Somerset, part l+II. Palaeonto- 
graphical Society Monographs 119: 1-109. 
1972. Permian gastropods and chitons from Perak, 
Malaysia. Part I, Chitons, bellerophontids, 
euomphalids and pleurotomarians. Bulletin of 
the American Museum of Natural History 147 
(1): 5-44. 

1985. Permian gastropods from Perak, Malaysia. 
Part 3, The murchisoniids, ceritiiiids, loxonema- 
tids, and subulitids. American Museum 
Novitatcs 2829: 1-62. 
BLODGETT, R.B. 1992. Taxonomy and paleobio- 
geographic affmities of an early Middle Devonian 
(Eifelian) gastropod faunule from the Livengood 
Quadrangle, east-central Alaska. Palaeonto- 
graphica, Abteilung A: 221: 125-168. 

1993. Dutrochus, a new microdomatid 
(Gastropoda) genus from the Middle Devonian 
(Eifelian) of west-central Alaska. Journal of 
Paleontology 62(2): 194-197. 
BLODGETT, R.B. & JOHNSON, J.G 1992. Early 
Middle Devonian (Eifelian) gastropods of central 
Nevada. Palaeontographica, Abteilung A:222: 83-139. 

1995. Merrianiites, a new name for the bel- 
lerophontid gastropod genus Merriamella 
Blodgett and Johnson. Journal of Paleontologv 
69(3); 608. 

BLODGETT, R.B. & ROHR, D.M. 1989. Two new 
Devonian spine-bearing pleurotomariacean 



CiIVKTIAN GASTROPODS FROM CiF.RMANY 57J 



gastropod genera from Alotk^,- l{^()T]dbtL'k>r 
Paieowology 63( I k 47-52, 
BLODGETT, R.B„ ROHR, & BGtJCOT* A.I. 
198S. Lq(v^ Devonian gsstcapoi biogcograDhy 
x^ttr W^vt} Hetpi^here^ In Embry, Ma^ il tan 

the StCQUd TfltemtlohftI syYtifpotitiRi the 

Devonian System: VoUrmc fll, Paleontolo^, 
paleoccolugy and biosiraiigraphy. Memoir 
Otnaclian SoQ^ PetrgJeuin Geolisgisis Mr 
281-294. 

1990. Early ;ind Middle Devonian gastropod bio- 
gcogmphv. VlL-nuiii (icoK»gti.;il SociclN ol 
London 277-284, 

CtlAPMAN» F, 1910, New or litllL^-lvnown X'iclorian 
fossils in the Naiional Museum. I'arl XIX. The 
Yeriniiian guxtrop(>d fauna. Proceedings gf \hf^ 
RoyarSocict>^of Vict0riav29 (0:75-103* 

COOK. A G. Flelchetvtevaio septata: a new 
high-spjred, septate gastropod iixm the Devonian 
of Nortki Qiteenstaod. Journal qf Folcointology 

!997. Gastropods from the Burdekln Ftirmatton, 
Middle Devonian. North Queensland. Memoirs 

of The Oiit-'ensiand Museum 42(1 ), 77-^!'> 
COOK, A G CAMHJ.ERL N. M»7 Middle 

l.>ev<.»maii yuslropodN Ironi the [Hrokeii Ki\er 

Pn.'vince. North Queensland Memotrs uf the 

Qut-enhland Museum 42(1 ): 55^79. 
DONAI f"). I IS91 Notes on some new and littlc- 

kn<»\vn spL-cif?: ot* C 'drhoniterous Miirc/u'sunja. 

<,>uai'lcrK Journal of the Geological Slocict>' of 

London: 562-575. 
J 895. Notes on the ^cnusMurchisomusmd its allies: 

with a revision of the Bfilisft CarhonifenJUs 

Species, eaad iiGScriptions of someMipw^ fbiins'. 

Quarterly journal of the Geological "Soctiisty tiC' 

London 51: 210-234. 
FR^DA. J. 1992. Mode of life of a new onychoctiSlid 

mollusc from the Lower Oevoni:iri oT HohcWjA. 

Journal of Paleontology ^(.(2)- 200-205 

Oldcsi |■eple:^cnloIivc of the tjinily P;n|noo- 

/.vgupleundac ICiaslrupodaJ with notes on ils 

hlglicr toxoDQiny. JounmT^PDJco9itology 4i7(5}; 

822-^27. 

W98a. Classilicafion and Phylogeny ot DiNoman 
Gastropods. UapubL thesis. Universitat 
Hnmburg, Hanlfeg«tgw 

1998b. Did the an^lors of higher gastropods 
(Neritimoiphay Caenogastropod^ and Kdeixi* 
sSxppta) smnfidjled shell? Abstracts, l3t& 
fnterhatlortal M^lHcological Congress,' 
Washington DC: 107. 

1998e. Some new and better recognized Dc\^onian 
gastropods from the IVague Basin (Bohemia). 
Bulletin of the C^cch Geological Survev 730): 
41-4Q. 

I9*^8d. Some new and heilL-r iccognized Devonian 
gastropods from ihc IVajrue Uasin - Bohcnua): 
part IL BuUeiin of the Czech Geological Survey 



4^99^j. riiiee new gastropod genera from Ihc Larly 
Devonian of 3oi)jQn)ia. dultoti^ of \ht Cjoach 

i999b. Higher ciassfncatfoii ^ Jutetfai^Jc 
gadtropods inferted fVom thBir Wly 5Ik«|] 
ontogeny hiurnal of the Czech Geological 
Society, Bairande volume. 44(1 -2): 137-153. 

PRtDAj.A BANDLL.K. 1097. New Early Devonian 
gaslro|n»ds from the PkrltrntVus (Boncoiomtim.!^ - 
Pi3lat'<r.\\;iii)U-uro ( '(miinunii) in the Prague 
Basin Bohemia) Mliteilungen aus dcni 
Cie(»h);^isch-I*alaconlolo£:isclien Itistjtui dtr 
Univcriviim l lamburuSO: 1-58. 

FRVDA, J. & MANDA, S. TW. A gastropod fawnule 
from the Kfonograpws unffbrmis graptolilc 
Hio/uiie bail) Luchkovtan, Early Devonjan) in 
Bah<pllW. Miiieilun|5!«U0"& ^tei pf9jogiscli* 
Talacoiftologlscben lastiiut der UtiiversrtKt 
HamhuT>i.«0: 59-122. 

GOLDFUSS. A. 1844. Peitcfacta Gofmauiae, 3. 
MJuNsckiorO. 

GORDON. M. & S OtHbLSON. h.L. 1987. Late 
Mississippiiin Gastropods ot the Chainirtfin Shale, 
wc.st-eciilral t'lali. I nilcd Slates- Geological 
Survey Profcs:Monal Paper I lOH; Ml?. 

GU8ANOV, A.R, BLODGETT. K.B.& LI TOCHKIN, 
V.M. J995. Early Devonian (Pragian) gastropods 

Taltantoldgyj 69; 431-440. 

IIOARfu R.D. & sniRGEON, M.T. 1978. The 
Pcnjisylvanian gastropod genera Cyciozyga nnd 
Helminthozyga and the classification of the 
Pscudozygopletiridae. Journnl ofPcileontology 52 

(4); 850-858. 

HORNY R. 1055. PalaeoTy^oplcurid.ic nov fani, 
tia^aropodal ze strcdoceskeho devonu. Sboinik 
Ustredniho listavu gcologiclveho oddil 
palconlolog3ck\ 21: 17-159. 
1992. New Lower Devonian Gastropoda and 
[ergomya (Mollusca) of Bohemia. Casopia 
Narodniho Moze?i v Praise, 1-4: 99-1 10. 
1 994. Shell morphology and ontogeny of the Lower 
Dcvofiian Mnrchisoniacean gastrc^pod 
Hniuziivyii miituroyjtihra (Murchisoniaeea). 
Casopis Narodniho Miiavi v Pra/e, M: 81 -S^, 

}]ORNY. R. U)Rr)\N. M I09J Silurian and 
Dcvtmian Gasiropoda of Koniaiua. CaSQpis 
Narociniho Mu/ea v !Va/r. 1-4:71-S5. 

KASL. I. *t NISHIDA. L 1080. A MiddJe Devonian 
gastropod faiinule h'om the Naka/iilo FonxiMioi^ 
of Kitakaivu. Northeast Japan. BuHctijiitf 'ftts 
NatiunuJ ScientieMu$eLtl|l'C12: 73-89. 
19E8. TVo new Mkfdlo Itevonian gftstwpofls fi^- 
the Nakayaio Formation of Ritakami, Northeast 
Japan. Saito Ho On Kai Museum Special 
Publication 2; 257-203. 

KlRCTfNFR. IIS, lOlS Mitteldevonische (.asiro- 
(HkIcii van Sr.icleiiich in dei Life!. Voi handlnn^jcn 
des N;Uijrhistori.schen Vcrciiiis dcr prcus.stschL*u 
RhcinlBndc uod West^l<m8 U: TS!^26] . 



374 



MEMOIRS OF THE QUEENSLAND MUSEUM 



KNIGHT, J.B. 1934. The gastropods of the St. Louis, 
Mo., Pennsylvanian outlier. The Euomphalidae 
and Platyceratidae. Journal of Paleonlolosy 8(2): 
139-166. 

1941. Paleozoic gastropod genotypes. Geological 

Society, Special Papers 32: 1-510. 
1945. Some new genera of Paleozoic Gastropoda. 

Journal of Paleontology 19(6): 573-587. 
1956. New families of Gastropoda. Journal of the 

Washington Academy of Sciences 46(2): 41-42. 
KNIGHT, J.B., COX, L.R., MYRA, A., BATTEN. 

R.L., YOCHELSON, E.L. & ROBERTSON, R. 

1960. Systematic descriptions Archaeogastropoda. 

Pp. 169-324. Treatise on Invertebrate Paleontology. 

(Geological Society of America, Inc. and 

University of Kansas Press: Lawrence, Kansas). 
KOKEN, E. 1889. Ober die Entwicklung der 

Gastropoden vom Cambrium bis zur Trias. Neues 

Jahrbuch fur Mineralogie, Geologic und 

Palaeontologie, 6: 305-484. 
1897. Die Leitfossilien, ein Handbuch fiir den 

Unterricht und das Bestimmen von Verstein- 

erungen. 1-848. Leipzig. 
KONINCK, E.G. d. 1881. Faune du Calcaire 

Carbonifere de la Belgique, Gasteropodes, 

Annates du Musee Royal d'Histoire Naturelle de 

Belgique, VI: 1-170. 
1 883. Faune du calcaire carbonifere de la Belgique 4 

partie, Gasteropodes suite et fm). Musee Royal 

d'Histoire Naturelle de Belgique Annales, Serie 

Paleontologique 8: 1-240. 
LINSLEY, R.M. 1968. Gastropods of the Middle 

Devonian Anderdon limestone. Bulletins of 

American Paleontology, 54(244): 333-465. 
LOTZ, H. 1900. Die Fauna des Massenkalks der 

LindenerMarkbei Giessen. 205-2 16. (Marburg). 
NUTZEL, A. 1997: Uber die Stammesgeschichte der 

Ptenoglossa (Gastropoda). Geowissenschaften. 

Abhandlungen E 26: 1-229. 
OEHLERT, D. & OEHLERT, R 1888. Descriptions de 

quelques especes devoniennes du departemenl de 

la Mayenne. Bulletin de la Societe d'Etudes 

Scientifiques d'Angers 1887: 65-120. 
PAECKELMANN, W. 1922. Der mitteldevonische 

Massenkalk des Bergischen Landes. 



Abhandlungen der Preussischen Geologischen 

Landesanstalt91: 1-112. 
PERNER, J. 1903, 1907, 1911. Gasteropodes. In J. 

Barrande, Systeme silurien du centre de la 

Boheme. IV: L390. (Prague). 
ROEMER, F. 1876. Lethaea palaeozoica. Atlas. 

(Hannover). 

ROLLINS, H.B., ELDREDGE, N. & SPIELER, J. 
1971. Gastropoda and Monoplacophora of the 
Solsville member Middle Devonian, Marcellus 
Fonnation) in the Chenango valley. New York 
State. Bulletin of the American Museum of 
Natural Histor\' 144(2): 133-170. 

SCHLOTHEIM, E.F. 1820. Die Petrefactenkunde auf 
ihrem jetzigen Standpunkte durch die 
Beschreibung seiner Samlung versteinerter und 
fossier Uben^este des Thier- und Ptlanzenreichs 
der Vorwelt erlautert. 1-436. (Gotha). 
1822. Nachtrage zur Petrefactekunde. 1-100. 
(Gotha). 

STAUFFER, C.R. 1 909. The Middle Devonian of Ohio. 

Bulletin of Geological Survey of Ohio, Fourth 

series 10: 160-194. 
VOSTOKOVA, V.A. 1961. Paleozoiskie gastropody 

Novoj Zemli i ostrova Vajgac. Sbornik statei po 

paleontologii i biostratigrafii. Leningrad. 5-25. 
1966. Devonian and Carboniferous gastropods 

from the Rudnyj Altaj Moutains. - Voprosy 

Paleontologii Tom V: 38-52. 
WENZ, W. 1939-1943. Gastropoda, Handbuch der 

Paiaeozoologie, 1-6: l-1639.(O.H.Schindewolf). 
WHIDBORNE, G.F. 1891. A monograph of the 

Devonian fauna of the south of England. Part III. 

The fauna of the limestones of Lummaton, 

Wolborough, Chircombe Bridge, and Chudleigh. 

Palaeontographical Society: 155-250. 
1892. A monograph of the Devonian fauna of the 

south of England. Part IV. The fauna of the 

limestones of Lummaton, Wolborougli, Chircombe 

Bridge, and Chudleigh. Palaeontographical 

Society: 251-344. 
YOCHELSON, E.L. 1953. Jedria, a new subgenus of 

Naticopsis. Journal of the Washington Academy 

of Sciences 43: 3-65. 



RELATIONSHIP BETWEEN MASS AND LENGTH IN AUSTRALL\N ELAPID 

SNAKES 



A.E. GREER AND R SHINE 

Greer, A.E. 8l Shine, R. 2000 06 30: Relationship between mass and length in Australian 
elapid snakes. Memoirs of the Queensland hduseum 45(2): 375-380. Brisbane. ISSN 0079-8835. 

Least squares regression analysis of log mass on both log snout-vent and log lotal length for 
individuals of each sex of 14 populations often species of Australian elapid snakes indicates 
that in the 37 most robust data sets isometry occurs in 21 cases, negative allometry in ten 
cases and positive allometry in six. Isometry seems to be the most common kind of allometry 
in 'colubroid--shaped snakes. There are no cases of ditTerent kinds of allometr>' between the 
sexes in any one species. However, in Austrelaps ramsayi both measures of length indicate 
that mass is relatively greater in males than in females over the middle and large end of the 
size range. The population of regression lines for log mass on log length for large diurnal, 
surface-active elapids are bounded by Austrelaps ramsayi on the heavy side and by 
Pseudonaja textilis on the light side. These extreme morphological differences may be 
related to the species' extreme ecological differences. The fonner species is a frog eating, 
live-bearing inhabitant of a cool environment with a short growing season, whereas the latter 
is a lizard, bird and mammal eating, egg-laying inhabitant of a warm environment with a 
longer growing season. □ Allometfy, elapids, mass, sexual dimorphism. 

A.E. Greer, Australian Museum. 6 College Street, Sydney 20] 0; R. Shine. Biological 
Sciences AOS, University of Sydney 2006. Australia: 26 November 1999. 



Mass is probably the single most important 
co-variate of an organism's other biological 
variables (McMaTion & Bonner, 1983; 
Schmidt-Nielsen, I984;Calder, 1996). Although 
there is a vast amount of raw data on mass for 
Australian vertebrates in the literature, 
unpublished theses and researchers' notes, little 
of this information has been collated and 
SLUTimarised. In this paper we analyse published 
and unpublished data on mass and length in ten 
species of Australian elapid snakes in order to 
determine the nature of the allometric relation- 
ships and whether sexual dimorphism exists. 

MATERIALS AND METHODS 

Data on mass (gm) and snout- vent and total 
length (mm) were gathered from the literature 
and our own notes (for origin of R. Shine's data 
and up-dated identifications, see Shine 1977 and 
1 989, respectively). All data were transformed to 
base 10 logs prior to analysis. 

The relationship between mass and length was 
analysed initially for each sex using least squares 
regression. The comparison between mass and 
each measure of length between the sexes was 
examined using analysis of covariance with 
length as the covariant and sex as the factor. 
Homogeneity of residual mean squares of the 
regressions was verified prior to the ANCOVA. 
All analysis was done usina SYSTAT 9.0 
software. The 0.05, 0.01, and 0.001 levels of 



significance are indicated by *, ** and ***, 
respectively. 

Mass data were rejected if the specimen was 
known to be gravid, and total length data were 
rejected if the specimen had part of the tail 
missing. 

RESULTS 

We found unanalysed data suitable for 
regression analyses of log mass on log length for 
14 populations of ten species of Australian elapid 
snakes. Data sufficient for a comparison of the 
regressions between the sexes of the same species 
were available for nine species (all except Suta 
suta which was only represented sufficiently by 
males). The results of these analyses are given in 
Table 1. We also include for the sake of 
completeness, but do not discuss, some basic 
regression statistics for Acatifhophis cmiarcticus 
which are the only previously published results 
with possible relevance for allometry in 
Australian elapids. 

The data are of variable quality for the purposes 
of our analysis. For example, we discount data 
for any sex based on fewer than ten specimens. 
We also discount the data for the female Notechis 
ater as they almost certainly include gravid 
individuals. The specimens were collected in 
period of 19 October to 15 February, and 
although no note was made of whether females 



376 



MEMOIRS OF THE QUEENSLAND MUSEUM 



appeared gravid ornot(Mirtschin& Bailey, 1990), 
the collection period coincides with the repro- 
ductive season (Shine, 1987b). We also discount 
the data for Austrelaps superbus from King 
Island due to the apparent rounding oITof both 
lengths and weights, the latter in some cases 
apparently to the nearest 50g (Feam, 1994). We 
also suspect that the relatively low r- values for 
female Notechis scuiatus from Melbourne may 
be indicative of an atypical variable (Wath^irow, 
1997, 1999). Finally, we note that the lack of 
small females in Hemiaspis sigrmta, may distort 
both the allometry and the comparison with 
males (data R. Shine). In the following discussion 
we put to one side these qualified data (indicated 
by itahcs in Table I) and focus on the remainder. 

DISCUSSION 

Methodologically, mass correlates with both 
snout-vent length and total length about equally 
well. In those cases where the r- values differ, 
snout-venl length has a higher value in three 
cases, and total length has a higher value in five 
cases. However, in no case does the difference 
exceed 0.01. Furthermore, both length measures 
gave the same indication of allometry in all eases 
except for Pseudechis porphyriacus from New 
England. For each sex in this species, the data for 
the total length indicated a relatively larger 
increase in mass with length than did snout-vent 
length. This could be due to a relatively larger 
increase in mass of the tail with length in this 
species. 

The relationship between mass and length in 
Australian elapids can be in positive aliometr>', 
isometry, or negative allometry, depending on 
species. However, isometry is the most common 
form of allometry, occurring in 21 of 37 cases 
('total samples' and 'combined' sexes excluded); 
negative allometry occurs in ten cases, and 
positive allometr>' occurs in six cases. In terms of 
the kind of allometry, there is no difference 
between the sexes of any species. Therefore we 
assimie that for species represented adequately 
by only one sex, the allometry of that sex is 
indicative for the species. 

In terms of the kind of allometry shown, the 
only comparable data set we are aware of is that 
for 12 species of North American colubrids, a 
group which is similar in shape and ecology to 
Australian elapids. The data for these species 
w ere based on combined sexes, but they indicated 
that the mass on length relationship was in 
isometry in 1 1 species and in negative allometry 
in only one, Heterodon platyrhinos (Kaufman & 



Gibbons, 1975). Hence it may be that isometry is 
widespread in 'cohibroid'-shaped snakes. 

Assuming that isometry is the null condition, 
the cases of negative and positive allometry in 
Australian snakes are of interest. Negative 
allometry occurs in Hemiaspis signata^ Notechis 
scutatiis from the New England area, and 
Parasuta dwyeri. Positive allometry occurs in 
Notechis scutatiis irom Melbourne, Pseudechis 
porphyriacus from Macquarie Marshes and, in 
tenns of total length only, in P. porphyriacus 
from New England. We have no explanation for 
these differences, but take note of the practical as 
well as the biological implications of the 
difference between two populations of the one 
species, Notechis scutatus. 

Differences between the sexes in the elevations 
in the slopes occur in three species: with both 
measures of length in Austrelaps ramsayi and in 
Notechis scutatus from the New England area, 
and in total length in Hemiaspis daemeli. How- 
ever, in the latter two species, the difference in 
elevations is so slight that we are reluctant to 
interpret it. In the case of Austrelaps ramsayi, it is 
clear on both measures of length that mass is 
relatively greater in males than in females over 
the middle and large end of the size range ( Fig. 1 ). 
This may be due to more, or denser, muscle mass 
as has recently been demonstrated in two 
colubrids and one viperid in Europe (Bonnet et 
al., 1998). 

The only species in which the residual mean 
squares of the regressions for each sex were 
significantly different was Pseudechis por- 
phyriacus from the New England area; the 
unexplained variance in mass was significantly 
higher in males than in females in both length 
measures (Table 1 ). Again we are unable to offer 
a biological explanation, but we note the 
statistical implications for other studies of sexual 
dimorphism in mass on length relationships. 

The mass on length regressions examined here 
form a distribution of roughly similar regression 
lines. Looking more closely al males only from 
the largest data sets (basically the Shine data) and 
restricting the analysis to snout-vent length for 
simplicity's sake, the boundaries of this dis- 
tribution of regression lines are set by Ausn^elaps 
ramsayi on the relatively heavy side and 
Pseudonaja textilis (total sample) on the 
relatively light side (Fig. 2). The regression lines 
of the males of Hemiaspis daemeli, H. signata, 
Notechis scutatus, Parasuta dwyeri and Pseudechis 
porphyriacus (not figured to avoid crowding) fall 



MASS AND LENGTH RELATIONSHIP IN- ELAP© SNAKES 



377 



3,2 




2.3 2.e 2.9 3.2 

p]GL L Plot tif ipgih^ log total length for hoih 
si^jissfXif^iixtFelc^ ramc^ ^om the New England 




FIG 2. Ploi of log JtiaSS ptirlog snoul-veni length fbv 
males of Aiistrelaps- rtmsayt ip) and ^seudam^a 



bctAvccn the regression lines lor Austrehips 
ramsayl and Psciuhnaja fcxfilis. In man\ ways 
these ianer two ta.\a represent the near extreme 
end$ of thetangccrf'diVerst^iDf^ti^^ 
surface dwelling Australian eiapids. For ex- 
ample, the former occurs at relatively high 
altitudes wilh shorlei. cooler gi'ovving seasons, 
feeds largely on tiogs atidis live bearing (Shine, 
1^^87a) whereas the latfer oectirs at lower -alti-^ 
tudes with a longer, warmer gniwing season, 
feeds largely on reptiles, birds and mammals and 
is egg laying (Shine, 10.S9). We do nol know 
what tissue differences may be contributing to the 
overaU diffejiBlices in mass in these two species. 
However, if it is fat» liiie heavi^ body in Ui^ 
species occupying the Cooler clithate with Ihfe 
shorter growing season makes adaptive sense. If 
it is muscle, the heavier body in the species 
hunting the getierally less acme prey and 
carrying. the lejiroductive Joad tor longer periods 
is-abo expected ftte rtauSstJtileJngtfr otflfw^^ 
in the females of the two species are almost, but 
not quite, as well separated as ni the males). We 
Stl^ecl that Denuitisia psanvnophis and perhaps 
Oxyiimms scutellauis would be even lighter for 
their length than is Psmfo^aja texHiiSi lujit we 
Idck the data to tesi this. 

This rev iew of mass and length data for Aus- 
tralian eiapids suggests to us several ways t(> 
improve the qualin- of these kinds ofdata. First, 
the two measures of length used, snoui-venl 
length an4 total l^gdi>^ ^cryit eqijaUy robm 



in tlieir reUitionship With mass. Hence although 
both measures arc usually taken, either will do by 
itself. Second, data on mass and length are most 
useful 5if Sfpecimens with potential confounding 
features are noted. Specilic*ally, specimens with 
missing tails, in obvious poor health, recently fed, 
or gra\id should have their condition indicated. 
Sexes should also be determined. Furthermore, 
HiiUs i^gdr fK^ sample size, tlie smaller the 
sampling area, and the more ponstr^in^d the col- 
lecting period, the tighter 'thd^ mass bn'tengtil 
relationship is likely to be Finally, it would 
veiy useful to have additional data for the smaller 
species of eiapids, as the CurTfiTrt dat^ Set tS 
primarily for large species. 

ACKNOWTEDGEMFNTS 

We thank J. Scanlon and G. Shea for critically 
readtflg eai4|r version^^of tbejn^^ 

LITERATURE CITED 

BONNET X., SHINE. R.. NAULLEAU. G & 
VACHERrW.LAS, M. 1998. Sexual dimorph- 
ic iij^nakes: dilTerent reproductive roles favour 
dimrgnt body plans. Proceedings of the Royat 
Society of London 265: 179-183, 

CALDER, W.A. 1996. Size, Ftmction, and Life Histoo'- 
(Dover Publications. Miiieola. New York). 

FEARN.S. 1994. Some obsciA -ations on the ecology of 
the copperhead Aiisn-elaps superhm (Serpcrites; 
Elapidac) in lasniania. 1 lerpctofauna 24: 1-10. 

n ARX.S c^c \1I1NDAY,B, 1995. The copperhead or 
superb snake in captivity. Monitor (Journal of tlie 
Wbori&ft Heipetolo^cal Soci^) 7: 9^Lf , 



378 



MEMOIRS OF THE QUEENSLAND MUSEUM 



JOHNSTON, GR. 1987. Reproduction and growth in 
captive death adders Acanthophis antarcticus 
(Squamata: Elapidae). Transactions of the Royal 
Society of South Australia 111: 123-125. 

KAUFMAN, GA. & GIBBONS, J.W. 1975. Weight- 
length relationships in thirteen species of snakes 
in the southeastern United States. Herpetologica 
31: 31-37. 

McMAHON,T.A.& BONNER, J.T 1983. On size and 
life. (Scientific American Books: New York). 

M1RTSCHIN,RJ.& BAILEY, N. 1990. A study of the 
Krefft's black tiger snake Notechis ater ater 
(Reptilia: Elapidae). South Australian Naturalist 
64: 52-61. 

SCHMIDT-NIELSEN, K. 1984. Scaling. Why is 

animal size so important? (Cambridge University 

Press: Cambridge). 
SHINE, R. 1977. Reproduction in Australian elapid 

snakes. II. Female reproductive cycles Aust. J. 

Zool. 25:655-666. 
1987a. Ecological ramifications of prey size: food 

habits and reproductive biology of Australian 



copperhead snakes {Austrelaps, Elapidae). Journal 
ofHerpetology 21(1): 21-28. 

1987b. Ecological comparisons of island and 
mainland populations of Australian tigersnakes 
(Notechis: Elapidae). Herpetologica 43: 233-240. 

1988. Food habits and reproductive biology of 
small Australian snakes of the genera Unechis 
and Siita (Elapidae). Journal of Herpetology 22: 
307-315. 

1989. Constraints, allometry, and adaptation: food 
habits and reproductive biology of Australian 
brown snakes {Pseudonaja: Elapidae). Herpet- 
ologica 45: 195-207. 

WATHAROW, S. 1997. Ecology of Eastern Tiger 
Snake (Notechis sciitatiis) and Lowland Copper- 
head (Austrelaps superbiis) within metropolitan 
Melbourne. Monitor (Journal of the Victorian 
Herpetological Society) 8: 145-151. 

1 999. Oophagy in the lowland copptxht^d Austrelaps 
siiperbus (Elapidae) in the Melbourne metro- 
politan area. Herpetofauna 29: 1 9-20. 



TABLE 1. Parameters for least squares regression analysis of logjQ weight (g) on logjQ length (mm) for ten 
species of Australian elapid snakes, ci =^ 95 percent confidence interval of the slope. A slope of 3.00 indicates 
isometry. All regressions are significant at the 0,001 level. 



Species 


Regression Parameter 


Comparison of Regressions (F values) 


inter | slope 


ci 




n 


allometry 


homogen 


slopes 


elevations 


Reference 


Snout-vent Length 


Austrelaps ramsayi New England 


males 


-5.92 


2.92 


±0.21 


0.93 


57 





1.32"^ 


0.96"' 


5.01* 


R. Shine data 


females 


-5.48 


2.74 


±0.28 


0.97 


13 













A. siiperbus Melbourne 


males 


-3.16 


J. 94 


0.66 


0.90 


6 




2.22"' 


4.95* 




Watharow, 
1997, 1999 


females 


-5.80 


2.86 


0.54 


0.93 


10 













A. superbus King Island 


males 


-2.98 


1.95 


0-40 


0.85 


20 




2.96"" 


13.34** 




Feam, 1994; 

Feam & 
Munday, 1995 


females 


-8.60 


3.82 


1.24 


0.84 


9 













Hemiaspis daemeli Macquarie Marshes 


males 


-6.20 


2.95 


±0.25 


0.96 


26 





2.38"' 


0.02"' 


0.81"" 


R. Shine data 


females 


-6.10 


2.92 


±0.40 


0.94 


15 













combined 


-6.15 


2.93 


±0.22 


0.95 


41 













H. signata New England area 


males 


-4.79 


2.39 


±0.26 


0.94 


24 




/.08^ 


0.12"" 


1.32"-' 


R. Shine data 


females 


-5.02 


2.49 


±0.57 


0.85 


17 













Notechis ater Flinders Ranges 


males 


'10.81 


2.43 


±0.57 


0.94 


9 





1.07^' 


3.3^ 


6.89*** 


Data in 
Mirtschin & 
Bailey, 1990 


females 


-13.89 


2.92 


±0.44 


0.95 


39 











Data in 
Mirtschin & 
Bailey, 1990 



MASS AND LENGTH RELATIONSHIP IN ELAPID SNAKES 



379 



TABLE L (co«0 



Species 


Regression Parameter 


Comparison of Regressions (F values) 


inter 


slope 


ci 




rt 


allometry 


homogen 


slopes 


elevations 


Reference 


Notechis scutatus Melbourne area 


males 


-7.39 


3.38 


±0.36 


0.94 


26 


+ 


1.34"s 


4.52* 




Data in 
Watharow, 1997 


females 


'5.29 


2.63 


±0.67 


0.79 


20 













N. scutatus New England area 


males 


-5.35 


2.66 


±0.15 


0.95 


73 




1.13"^ 


1.45"'^ 


4.42* 


R. Shine data 


females 


-5.66 


2.79 


±0.15 


0.97 


51 












Parasuta dwyeri New England area 


males 


-5.31 


2.63 


±0.23 


0.93 


39 




1.37"^ 


0.057"^ 


1.28"' 


R. Shine data 


females 


-5.17 


2.58 


±0.31 


0.91 


31 












combined 


-5.17 


2.58 


±0.18 


0.93 


70 












Pseudechis porphynacus Macquarie Marshes 


males 


-7.04 


3.25 


0.17 


0.98 


37 


+ 


}.4r 


12.08** 




R. Shine data 


females 


-3.35 


2.02 


0.58 


0.91 


7 












P. porphyriacus New England area 


males 


-6.68 


3.12 


0.20 


0.89 


128 











R. Shine data 


females 


-6.67 


3.11 


0.19 


0.96 


49 













Pseudonaja textilis Macquarie Marshes 


males 


-6.15 


2.89 


±0.52 


0.83 


27 





7.05'" 


1.2(r 


0.004"" 


R. Shine data 


females 


-7.78 


3.42 


±0.80 


0.91 


9 













combined 


-6.17 


2.90 


±0.18 


0.97 


37 













P. textilis New England area 


males 


-6.54 


3.02 


±0.14 


0.99 


18 





1.36"^ 


0.05"' 


0.83"' 


R. Shine data 


females 


-6.49 


3.00 


±0.18 


0.99 


11 













combined 


-6.56 


3.03 


±0.10 


0.99 


29 













P. textilis Total Sample 


males 


-6.46 


2.99 


±0.14 


0.98 


45 





1.09"^ 


0.05"' 


0.32"" 


R. Shine data 


females 


-6.54 


3.01 


±0.15 


0.99 


20 













combined 


-6.52 


3.01 


±0.10 


0.99 


65 













Suta suta New South Wales 


males 


-5.90 


2.83 


±0.24 


0.98 


14 











A. Greer data 
and Shine, 1988 


Total Length 


Acanthophis antarcticus Eyre Peninsula 


combined 


-6.76 


3.09 


? 


0.97 


206 


? 


? 


? 


7 


Johnston, 1987 


Austrelaps ramsayx New England area 


males 


-6.42 


3.02 


±0.19 


0.95 


54 





l.Ol"" 


223"^ 


5.09* 


R. Shine data 


females 


-5.76 


2.76 


±0.29 


0.97 


13 













Austrelaps superhus Melbourne area 


males 


-3.24 


1.93 


0.68 


0.89 


6 




i.3r 


2.59^' 


i.o(r 


Watharow, 
1997, 1999 


females 


-5.80 


2.86 


0.54 


0.93 


10 













A. superbus King Island 


males 


-3.72 


2.14 


0.44 


0.84 


20 




3.08^ 


11.22** 




Feam, 1994; 

Feam & 
Munday, 1995 


females 


-9.44 


3.99 


1.34 


0.83 


9 













Hemiaspis daemeli Macquarie Marshes 


males 


-6.54 


2.99 


±0.25 


0.96 


26 





2.00"' 


0.0 r 


6.25* 


R. Shine data 


females 


-6.41 


2.97 


±0.36 


0.95 


15 














380 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE L (cont.) 



Species 


Regression Parameter 


Comparison of Regressions (F values) 


inter 


slope 


ci 


r^ 


n 


allometry 


homogen 


slopes 


elevations 


Reference 


Hemiaspis signata New England area 


males 


-5.20 


2.46 


±0.25 


0.95 


23 




1.12"" 


0.09^ 


4.26* 


R. Shine data 


females 


'5.39 


2.55 


±0.54 


0.86 


17 













Notechis ater Flinders Ranges 


males 


-11.62 


2.49 


±0.59 


0.93 


9 





l.KT 


2.54"" 


7.26* 


Data in 
Mirtschin & 
Bailey, 1990 


females 


-14.49 


2.95 


±0.24 


0.94 


38 













Notechis scutatus Melbourne area 


males 


-7.80 


3.43 


±0.29 


0.93 


46 


+ 


1.97"" 


6.47* 




Data in 
Watharow, 1997 


females 


-5.58 


2.67 


±0.72 


0.76 


19 













Notechis scutatus New England Area 


males 


-5.77 


2.1 A 


±0.14 


0.96 


70 




1.25"' 


0.06"' 


5.71* 


R. Shine data 


females 


-5.80 


2.76 


±0.15 


0.97 


48 












Parasuta dwyeri New England area 


males 


-5.51 


2.65 


±0.23 


0.93 


39 




2.07* 






R. Shine data 


females 


-5.30 


2.58 


±0.31 


0.91 


31 












combined 


-5.25 


2.56 


±0.18 


0.93 


70 












Pseudechis j 


vorphyriacus Macquarie Marshes 


males 


-7.52 


3.33 


±0.17 


0.98 


36 


+ 


1.33"" 


11.66** 




R. Shine data 


females 


-3.75 


2.10 


±0.62 


0.92 


6 












P. porphyriacus New England 


males 


-7.20 


3.23 


±0.20 


0.89 


117 


+ 


3.51*** 






R. Shine data 


females 


-7.20 


3.22 


±0.20 


0.96 


44 


+ 










Pseudonaja textilis Macquarie Marshes 


males 


-7.05 


3.10 


±0.65 


0.80 


25 





1.15'^ 


0.43"" 


0.0^ 


R. Shine data 


females 


-8.21 


3.47 


±0M 


0.90 


9 













combined 


-6.49 


2.92 


±0.20 


0.96 


35 













P. textilis New England 


males 


-6.66 


2.98 


±0.10 


0.99 


16 





3.11"'' 


0.0 19"" 


0.52"^ 


R. Shine data 


females 


-6.72 


3.00 


±0.21 


0.99 


10 













combined 


-6.71 


3.00 


±0.10 


0.99 


26 













P. textilis Total sample 


males 


-6.64 


2.97 


±0.13 


0.98 


41 





1.37"" 


0.05"' 


0.59"" 


R. Shine data 


females 


-6.72 


2.99 


±0.16 


0.99 


19 













combined 


-6.70 


2.99 


±0.10 


0.98 


60 













Suta suta New South Wales 


males 


-5.98 


2.79 


±0.24 


0.98 


14 











A. Greer data 



FRESHWATER DALYELLIID FLATWORM, GIEYSZTORIA SUPERBA SR NOV. 
(DALYELLIIDAE: RHABDOCOELA) FROM SOUTHEAST QUEENSLAND, 

AUSTRALIA 



V. HARTENSTETN AND K.A. DWINE 

Hartenstein, V. & Dwine, KA. 2000 06 30: Freshwater dalyelliid flatworm, Gieysztoria 
superha sp. nov. (Dalyelliidae: Rhabdocoeia) from southeast Queensland, Australia. 
Memoirs of the Queensland Museum 45(2): 381-383. Brisbane. ISSN 0079-8835. 

A new dalyelliid flatworm, Gieysztoria superha sp. nov. is described from a freshwater creek 
in southeast Queensland, Australia. With characters of the genus, it appears to be closely 
related to the cosmopolitan species G rubra, but differs significantly in the number of spines 
(approx. 40) in the sclerotic armature of the male copulatory organ. □ Freshwater, 
Dalyelliidae, taxonomy Australia. 

V. Hartenstein, Department of Molecular. Cell and Developmental Biology, UC Los 
Angeles, PO Box 951 606, Los Angeles CA 90095-1606, USA; K.A. Dwine, Queensland 
Museum, PO Box 3300, South Brisbane 4101. Australia: 26 November 1999. 



The Dalyelliidae contains many, mostly 
free-living, freshwater species of flatworms that 
are known from around the world ( Luther, 1955), 
and in Australia ultrastructural studies have been 
conducted on an unidentified species of the cos- 
mopolitan aenus Gieysztoria by Rohde, Cannon 
& Watson (1988) and Watson & Rohde (1995). 
No taxonomic study of the family or its members, 
however, has been conducted here. 

METHODS 

Freshwater plants and sediments were collect- 
ed from Gold Creek, Brooktleld (27°27^S8"S; 
1 52°53 '03"E), reuimed to the laboratory, and the 
worms extracted by carefijl examination using a 
stereo microscope. Worms were examined alive 
with bright lield and Nomarski interference con- 
trast microscopy. Whole mounts in Canada 
balsam were made after specimens were fixed in 
5% buffered fonnalin and dehydrated through a 
graded alcohol series. Specimens were not 
stained. Sclerotic armature preparations were 
made by squash preparation and mounting in 
Epon or by immersing live worms in de Faure's 
lluid (Evans et al., 1961). Sections (2|jm) were 
obtained serially from worms fixed in 4% 
glutaraldehyde in phosphate buffered saline and 
embedded in Epon. Sections were stained with 
toluidine blue. 

Drawings of the anatomy were made using a 
camera lucida and digitised using Adobe 
Illustrator®. Measurements are in micrometres 
([iva) unless otherwise stated. 

This material is lodged at the Queensland 
Museum (QM) as a wholemount (WTVI), serial 
sections (TS) and squash preparations (S). 



DALYELLIIDAE Graff, 1908 

Gieysztoria superba sp. nov. 
(Figs 1-2) 

MATERIAL. HOLOT\TE: Gold Ck, Brookfield SEQ 
(27°27'58"S, 152°53'03"E). Feb. to Apr. 1998, V. 
Hartenstein, WM, QMG2 17989. PARATYPES: same 
data, TS (Epon: toluidine blue), _QMG2 17990. OTHER 
MATERIAL: same data. Sclerotic amiature preparation 
(squash preparation, Epon) QMG2 17991. 

ETYMOLOGY. Named superba for the large number of 
spines in the male copulatoiy amiature. 

DESCRIPTION. HOLOTYPE: 550 long x 200 
wide (Fig. 1), clear, filled with brown algal? 
bodies and posteriorly red pigment bands. Swims 
swiftly over substrate; frequently rotates around 
length axis while swimming. Anatomically (from 
live material and sections) consistent with the 
genus and having finger-like vitellaria. Arising 
from the distal portion of the muscular bulb 
surrounding the ejaculatory duct and prostate 
secretions there is a fenestrated girdle (20 wide) 
with irregular holes (Fig. 2). Suspended distally 
from the girdle are about 40 fine spines of about 
equal length, evenly arranged around the whole 
duct. Each spine is fluted with a fine tip and 
approx. 43 long. 

DIAGNOSIS. With characters of the genus and a 
male copulatory organ with a moderately narrow, 
fenestrated girdle and about 40 spines of equal 
length. 

HABITAT AND DISTRIBUTION. Found in the 
sediments and plants of freshwater creek at 



382 MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1 . Gieysztoria siiperha sp. nov. Habitus. Scale = 
200^m. 

Brookfield, SE Queensland. Numbers appear to 
decline in the colder months (May-September). 

REMARKS. Luther (1955) divided the 46 
species of Gieysztoria then described into two 
groups: 'aequales' with a sclerotic armature 
displaying a wreath of homogeneous spines 
(about 25 species) and 'inaequales' forthe rest. G 
super ba belongs to the first group: it has a 
sclerotic armature with a girdle and possesses 
papillose vitellaria. In the numerous spines G 
superba resembles G knipovici Beklemischev 




FIG. 2. Gieysztoria superba sp. nov.; two focal planes 
of the sclerotic armature in a squash preparation (QM 
G2 17991). Scale = 50^011. 

1 953 which has at least 30 spines in its armature, 
but G superba differs in having vitellaria with 
long, digitate extensions (not short papillae). G 
superba has similar vitellaria to G. beUis Marcus 
1946 and G. rubra Fuhrmann 1894. G. superba 
can be distinguished because its spines lack the 
cone-shaped tip found in G bellis. and although 
the armature of G. superba looks very similar to 
that of G rubra, it has a wider girdle and shorter 
spines. Most characteristic of G superba is the 
number of spines in its armature. While G beilis 
and G rubra have approx. 19 spines, G superba 
has approx. 40. 

DISCUSSION. The inter-specific variation in 
spine number between G superba and G rubra is 



A NEW FRESHWATER DALYELLIID FLATWORM 



383 



far greater than the intra-specific variation within 
G, rubra that is associated with geographic 
location. (Marcus, 1946; Luther, 1955; Young, 
1977). We, therefore, have no hesitation in con- 
sidering tliis a new species, the first taxonomic 
account of any member of tiie DalyeJliidae from 
Australia. 

ACKNOWLEDGEMEl^TS 

We thank Amelia llarlcnstein for her hcip in 
the field and Jaborator> and Lester Cannon fijr 
comments on the MS. Without the support of the 
NIH grant NS 29367 for VH this work would not 
have been initiated. The study was part of a 
broader study by KD supported on a grant to Dr 
L.R.G Cannon from AF-iRS. 

LITERATURE CITED 

EVANS, GO., SHEALS, J.G & MACFARLANE. 
196L The terrestrial Acari of the British Isles- an 
introductiou to their inQxphol<^ biolo^ and 
classiCcatton. Vol 1. Introductkn and ikH^, 
(Adlard & Soii» Bartholomew Press: Dorking, 
GreaA Britain). 



LUTHER, A. 1955, Die Dalyelliiden (Turbellaria 
Neorhabdocoela), elne monographie. Acta 
ZoologicaFennicaS?: 1-337. 

MARCUS, E. 1946. Sobre Turbellaria Brasil«ros. 

Zoologia 11: 5-254. 

ROHDE, K., CANNON, L.R.G & WATSON, N. l988. 
Ultrastructure of the flame bulbs and proto- 
ncphridial capillaries of Gieysztor'ia sp. 
(Rhahdocoela Dalyelliida), Rhinolasius sp. 
(Khabdocoela Kalyplorhynchia) and Aciino- 
daclylellu blanchardi (Rbabdocoela 
Temnocephalida). Journal of Submicroscopic 
Cytology and Pathology 20(3): 605-612. 

WATSON, N. & ROHDE, K. 1995. Ultrastructure of 
s p e rni i L) g e n e s i s and s p e rni a I o z o a in I h e 
plaly helminths AclinodaciylcUa blanvhanli 
(Temnocephalida, Actinodactylellidae). Didy- 
morchis sp. (Temnocephalida. Did\ morchidae) 
and Gieysztoria sp. (DalyeUiida, Dalyelliidac), 
with implications for the phylogeny of the 
Rhabdocoeia. Invertebrate Reproduction and 
Development 27(2):145-158. 

YOUNG, J,0, 1 977, Six new species and records of two 
established species of Dalyelliidae {Turbellaria; 
NeoiiiabdDCoela) from freshwater habitats in 
Kenya, East Africa. Journal ofNatural Hstory 1 1 r 
1-15. 



384 



MEMOIRS OF THE QUEENSLAND MUSEUM 



NEMATODES FROM FISH FROM THE KEPPEL 
ISLANDS, KEPPEL BAY, QUEENSLAND. Memoirs of 
the Queensland Museum 45(2): 384. 2000:- Seventy six 
fishes comprising 24 species collected (by members of the 
Capricorn Scuba Club) from three sites; the waters off Ross 
Reef (23°03'S, I50°22'E), Barron Island (23°08'S, 
15r03'E) and North Keppel Island {23°04'S, I50°53'E)in 
Keppel Bay on the central Queensland coast, were examined 
for nematodes. Thirteen species of nematode, all new locality 
records, were found (Table 1 ), No nematodes were found in 
Choerodon venustus De Vis or Acanthurus dussimeri 
Valenciennes and only unidentifiable larvae in Siganus 
Uneatus (Valenciennes). Names of the fish hosts follow 
Randell, et. al. (1990). Ascaridoid larvae were identified 
following Cannon (1977) but citing Thynnascaris as a 
synonym of Hysterothylacium (Deardorff & Overstreet, 
1981 ). Comparisons between the fish examined in this study 
and from Heron Island (Lester & Sewell, 1989) showed 
differences in the nematode communities. Of five nematode 
species recorded from Heron Island two, Philometra sp. and 
Metabronema magna, were not present in Keppel Bay. 



Cucullaniis sp. and Hysterothylacuim sp. larvae were found in 
C. venustus at Heron Island but not Keppel Bay, while 
Terranova sp. larvae was found only in Epinephalus ongus at 
Heron Island. 

Literature Cited 

CANNON. L.R.G, 1977. Some larval ascaridoids from south-eastern 
Queensland. International Journal for Parasitology 7: 233-243. 

DEARDORFF, T.L. & OVERSTREET, R.M. 1981. Review of 
Hysterothylacium and Iheringascaris (both previously ^ 
Thynnascaris) (Nematoda: Anisakidae) from the Northern 
Gulf of Mexico. Proceedings of the Biological Society of 
Washinslon 93: 1035-1079. 

LESTER. R.J.G. & Sewell, K.B. 1989. Checklist of parasites from 
Heron Island, Great Barrier Reef Australian Journal of 
Zoology 37: 102-128. 

RANDALL, J.E., ALLEN, G.R. & STEENE, R.C. 1990. The 
complete divers and fisherman's guide to fishes of the Great 
Barrier Reef and Coral Sea. (Crawford House Press: Bathursl). 

L.R. Smales, School of Biological and Environmental 
Sciences. Central Queensland University Rockhampton 
4702, Australia: 14 February 2000. 



TABLE 1. Nematode parasites found in 24 species offish collected in Keppel Bay. September 1989 to July 1990. b = body 
wall, in = intestine, li = liver, me = mesentery, st = stomach. 



Parasite Taxon 


Site 


Host Species 


Parasite Taxon 


Site 


Host Species 


Trichuridae 


cortt. 




Kvphosiis sp. 


CapiUaria sp. 


St. 


Caranx ignobilis (Forsskfil) 






P. batavianus 


in. 


Diaeramma picium (Thunbere) 






P. tiera 




Ephinephalus onsus (Bloch) 






PI. schotaf 






E. taiivina (Torsskil) 






Plectropomus ieopardus fLacepede) 






Kvphosits sp. 


Terranova sp. 


in. 


C ignobilis 






Plectorhinchiis flavomaculatus fCuvier) 


larva type 1 \ 


me. 


Choerodon schoenleinni (Valenciennes) 






PI. sibbosus (Lacepede) 




Cr. altivelis 






PI. schoiqf (ForsskSl) 






D. pictum 


Cucullanidae 






E. cvanopodus 


Cucullanus australiensis 


in. 


C isnohilis 






E. ongus 


Bay lis 1927 


D. inctum 






E. lauvina 






PI. scholaf 






Kvphosus sp. 


Dichelym (Dichelyne) sp. 


in. 


PI. gibhosus 






Lutjanus carpanatatus (Richardson) 


Anisakidae 






L russelli (Blecker) 


Anisakis sp. larva type 1 


li. 


PI. schotaf 






Naso hcxacanthus (Bleeker) 




D. pictum 






N. unicornis (Forssk^l) 


Contracaecum sp. 
larva type 1 1 


in. 


C. ignobilis 






P. teira 


Hysterothylacium sp. 
larva type 1 1 




E. cvanopodus Richardson 






P. batavianus 




PI. flavomaculatus 






PI. flavomaculatus 


Hysterothylacium sp. 


me. 


Cromileptes allivelis (Valenciennes) 






Pi gibbosus 


larva type 1 1 1 


in. 


E. cvanopodus 






PL schotaf 




E. lauvina 






S. sordidus (ForrskSl) 






Hemievmnus melapteriis (Block) 


Camallanidae 






Kyphosus sp. 


Procamallanus lonis Cribb, 
1989 




P. batavianus 






Pi flavomaculatus 


Camatlanus sp. 


St. 


Pi flavomaculatus 






PI. gibhosus 


Cysticolidae 






PI. scholaf 


Ascarophis sp. 


St. 


D. pictum 






Platax batavianus (Cuvier) 


E. cyanopodis 






P. teira (Forsskdl) 






E. lauvina 






Scarus rivulatus 






N. unicornis 


Hysterothylacium sp. 


me. 


C. ignobilis 






Pi gibbosus 


larva type IV 


in. 


D. picium 


Crislilectus sp. larvae 


in. 


Pi scholaf 




St. 


H. melapterus 


N. unicornis 



ANEW SPECIES OF STREPTOCEPHALUS (PARASTREPTOCEPHALUS) 
(CRUSTACEA: ANOSTRACA: STREPTOCEPHALIDAE) FROM NORTH 
QUEENSLAND, AUSTR^^LIA 



B. HERBERT AND B.V. TIMMS 

Herbert, B. & Timms, BA^ 2000 06 30: A new species of Streptocephalits 
(Parastreptocephalus) (Crustacea: Anostraca: Streptoccphalidae) from North Queensland, 
Australia. Memoirs ofthe Queensland Museum 4S{2): 385-390. Brisbane. ISSN 0079-8835. 

Streptocepholus (Parastreptocephalus) queenslandicus sp. nov. is described from temporar>' 
ponds near Atherton, north Queensland. It is related to four species of the recently erected 
subgenus Parastreptocephalus from Africa mainly because of male antenna! morphologv' 
and the production of tetrahedrai eggs, a unique feature within the family Streptoccphalidae. 
This confirms the presence of Streptocephalus in Australia in a continent whose fairy shrimp 
fauna is dominated by Branchinella and Parartemia. □ Parastreptocephalus, new species, 
Streptocephalus archeri, control in fish ponds, biogeography, fairy-shrimp. 

B. Herbert, Department of Primary Industries, Kennedy Highway. Walkamin. 4872; B. V. 
Timms, Department of Geography and Environmental Science, University of Newcastle, 
Callaghan, 2308, Australia; 24 May 1999. 



Despite the early description by Sars (1896) of 
Streptocephalus archeri from the Rockhampton 
area of central Queensland, and a further 
collection of six females from the same area 
(Linder, 1941), no other specimens of the 
Streptocephalidae have been found in Australia. 
Adding to the enigma, the description was based 
entirely on females, so that it is difficult to 
determine the relationships of S. archeri within 
the genus. It is not surprising then that subsequent 
studies on Streptocephalus have reported little on 
S. archeri (see Brendonck et al., 1992; Belk & 
Brtek, 1 995) and reviews of aquatic invertebrates 
in Australia have discounted the presence of 
Streptocephalus in Australia (Geddes, 1981, 
1983; Williams, 1980, 1981). 

Over the last few years fairy shrimps have been 
encountered by one of us (BH) each time 
aquacukure ponds were filled at the Freshwater 
Fisheries and Aquaculture Centre, Walkamin, on 
the Atherton Tablelands in north Queensland. 
Specimens collected in February 1 997 were ident- 
ified by BVT as a species of Streptocephalus. 
Further collections in October 1997 confirmed 
their presence and provided data on development 
times. The males are different to any known 
species of Streptocephalus and the females 
appear different from those ofS. archeri, so they 
are described here as a new species. 



CRUSTACEA 
ANOSTRACA 
STREPTOCEPHALIDAE Daday, 1910 
Streptocephalus Baird, 1852 
Parastreptocephalus Brendonck, Hamer & 
Thiery, 1992 

Steptocephalus (Parastreptocephalus) 
queenslandicus sp. nov. 
(Figs 1-3) 

ETYMOLOGY. From Queensland, where the specimens 
were found. This complements names based on localities 
in Africa tor many other species of the Parastreptocephalus 
(Brendonck et al, 1992). 

MATERIAL. HOLOTYPE: c^, QMW24520. PARA- 
TYPES: 9 ds, QMW24521; 10 9 's, QMW24522. 
Queensland Museum. Lengths: 6 holotype and paratypes 
12.1 ± 1.2mm; 9 paratypes 12.1 ± 0.9mm. OTHER 
MATERIAL: 46 adults collected from rearing ponds at the. 
Walkamin Research Station, Walkamin, via Atherton, N 
Qld, Feb. 1997, B. Herbert, QMW24579 Queensland 
Museum. 100+ inimature specimens from a subsequent 
filling of the same ponds collected by B. Herbert on 1 6 Oct. 
1997, QMW24580, Queensland Museum. 

TYPE LOCALITY. Fish rearing ponds, Walka- 
min Research Station, Walkamin, via Atherton, N 
Qld, 17°8'S, 145°26'E, altitude 590m a.s.l. 
Collector B. Herbert, Feb. 1997 2-3 weeks after 
flooding of ponds. 

DESCRIPTION. Male. Antennules simple and 
long, almost reaching end of antennal base, 
tapering slowly towards a blunt tip which bears 
three setae (Fig. IE). Lateral processes of 
antennae smooth, slightly curved backward and 



386 



MEMOIRS OF THE QUEENSLAND MUSEUM 



1 mm 





1 mm 



FIG, 1. Streptocephalus (Parastreptocephalus) queens I and icu sp. nov., 
A, dorsal view of finger of antenna; B, inner or medial view of right hand of 
antenna; C, lateral view of right hand of antenna; D, dorsal view of frontal 
organ; E, tip of antennule; F, base of antenna. 



inserted at posteriobackward margin of distal end 
of basal segment (Fig. 1C,F). Median process of 
antenna continuing at same thickness of basal 
segment consisting of a short, slightly bent 
middle part and a terminal hand (Fig IB,C ). Basal 
part of hand about twice as wide as middle and 
whole hand about twice the length of the middle 
part. Much of the outside of the hand with a warty 
surface, as opposed to a smooth surface on the 
middle part and inner surface of the thumb and 
finger. Basal part of thumb grooved, with 
chitinised wavy edges (Fig 1 B,C). The outer base 
carrying a curved pointed spur having one margin 
continuous with the chitinised wavy edge of the 



basal groove of the thumb 
(Fig. 1B,C). Distal half 
(anterior) of thumb attached at 
an obtuse angle (140-160°) 
and with a basal protuberance 
marking the end of the groove 
(Fig. 1B,C). Distal end of the 
thumb tapering to a slightly 
recurved point reaching well 
beyond the tip of the finger. 
Finger with two teeth, one on 
the inner margin and the other 
on the upper surface, so that 
the spur of the thumb extends 
near both (Fig. lA-C). The 
upper marginal tooth recurved 
and with a blunt point, while 
the tooth on the inner surface 
is rounded (Fig. lA). Distal to 
the rounded tooth is a smooth 
groove tlanked on the inner 
margin with a warty 
protuberance near the tooth 
and a warty surface elsewhere 
(Fig. 1 A). Apical part of tooth 
curved backward. Frontal 
appendage small with a 
double blunt point (Fig. ID). 

Thoracopods with five 
endites, decreasing markedly 
in size distally. Many long 
posterior setae, but with a 
restricted number of anterior 
setae, as itemised in Table 1 
and shown in Fig. 2A-F. 
Endopodite broadly truncated 
with an apical shallow notch 
medially. External margin of 
endopodite with hook-like 
setae, but medially, especially 
towards the medial-distal 
comer setae longer, straighter and plumose (Fig. 
2G,H). Base of endopodite setae with 2-8, usually 
3-4, very small spines. Exopodite ladle-shaped 
and with numerous long plumose setae, but with 
a few small stout tooth-like setae at the base of the 
external margin (Fig. 2A,1,J). The long setae also 
attended at their base with very small spines, 2-6 
in number but usually 3-4. Epipodite rather 
similar in shape to the exopodite, but smaller and 
without marginal setae. Preepipodite lamelliform 
and with irregular small hooks on the external 
margin (Fig. 2A,K). 

First ten pairs of thoracopods similar except for 
two extra anterior setae on endite 5 of limb 1. 



0-1 mm 



NEW STREPTOCEPHALUS FROM NORTH QUEENSLAND 



387 



Occasionally there may be reductions in the 
number of long anterior setae of endites 3-5 of 
limbs 2-10. Eleventh pair of thoracopods 
reduced, mostly at the inner proximal comer, so 
that endites 1 and 2 much reduced, but other 
components only a little smaller and less setose 
than for thoracopods 1-10. Anterior setae on the 
endites of limb 1 1 much reduced and the number 
of posterior setae reduced, but still >10, on 
endites 1 and 2. 

Basal and nonrelractable parts of penes each 
bearing near its base a posteriorly curved median 
sausage-like outgrowth with 3-4 teeth on the 
proximal surface (Fig. 3A,B)- 

Cercopods (Fig. 3C) lanceolate and separately 
attached to posteriorlaleral margin of a short 
telson. Each bears plumose setae on its inner and 
outer margins and the lip. 

Female. Anlennules filamentous, slightly longer 
than antennae or of equal length. Tip with three 
setae as in male. Antennae foliaceous and oval 
with broadly rounded margins and no apex (Fig. 
3E). Slight notch sometimes on the niid-distal 
margin. Margins edged with small weak setae, 
with very few near the basal portion. Brood 
pouch cylindrical, elongate, extending to the end 
of the sixth abdominal segment in mature 
specimens, i.e. pouch is four segments long (Fig. 
3D). 

Immature eggs spherical, but mature shelled 
eggs tetrahedral (Fig. 3F). Comers somewhat 
rounded and hyaline; edges also hyaline and 
slightly thickened. Planar surfaces slightly 
convex and w ith a small raised area in the middle. 
Within the hyaline thickening on the edges and 
comers the embryo is round. 

Thoracic appendages and cercopods as in 
male. 

DIAGNOSIS. The present specimens easily lie 
within Streptocephalus because the antennae in 
the male have a medial outgrowth from the basal 
joint which temiinates in a cheliform structure 
(Brendonck, 1990). Because of the tetrahedral 
eggs of the female and the well developed spur of 
the thumb and the lack of teeth between this spur 
and the anterior (main) part of the thumb, they lie 
within the subgenus Paras treptocephal us as 
presently defined (Brendonck et al., 1992). 

Though the genera! stmcture of the antennae, 
thoracopods, cercopods and the nonretractable 
part of the penes of the male are very similar to 
those of the four spQc'ics of Streptocephalus so far 
asssigned to Parastreptocephalus (Brendonck et 
aJ., 1 992), none have the same antennae stmcture 



TABLE 1. Idealised arrangement of setae on the 5 
endites of male thoracopods of.S'. cpwenslandicits. 
First figure indicates number of anterior setae, 
second the number of posterior setae, m = many (> 
10). 



Thoracopod 


Endite 


1 


2 


3 


4 


5 


1 St pair 


3 + m 


2 +m 


2 + 3 


2 + 2 


3+2 


1st - lOth 
pair 


3 + m 


2 +m 


2 + 3 


2 + 2 


1 +2 


1 1 th pair 


I +m 


1+3 


2 +-2 


2 + 1 


1 + I 



or setation of the endites of the thoracopods as in 
S.(P). queemlandicus . The teeth of the antemial 
finger are more rounded, spaced further apart and 
in different planes than in the other three species 
oi Parastreptocephalus with teeth on the finger. 
The endite setae are very similar to those of S. (Pj 
sudanicus and S.(P.) zuluensis, the two species of 
Parastreptocephalus that have been studied in 
detail (Brendonck et al., 1992). The only con- 
sistent di (Terence is the presence of three instead 
of four anterior setae on the first endite in S. (P.) 
queemlandicus. 

Although S. archeriwas incompletely described 
by Sars, there are a number of differences between 
it and .S". queenslandicus. The medial distal part of 
the antenna has no short pointed projection in 5. 
queenslandicus as it has in S. archeri. The brood 
pouch is four segments long in S. queemlandicus 
whereas it is only two in S. archeri. Part of this 
difference may be explained by the greater 
overall length of S. queenslandicus (12mm as 
against 9mm for S. archeri). The tetrahedral eggs 
of S. archeri ^exhibit plane or very slightly 
concave faces connected by obtuse, prominent 
ribs' (Sars, 1896), though the diagrams show 
them w ith markedly concave planar surfaces. By 
contrast the eggs of S. queenslandicus have 
slightly convex surfaces and the ribs (i.e. the 
edges of each tetrahedral surface) are not obtuse 
or particularly prominent. S. archeri eggs, while 
tetrahedral, are more like those of S. sudanicus, 
while those of S. queenslandicus are more like 
those of S. zuluensis particularly in the rib 
structure, though the planar surfaces are slightly 
convex in S. queenslandicus and slightly concave 
in S. zuluensis (see Brendonck et al., 1992). 
These apparent differences in egg stmcture may 
be artefacts of preparation and so of limited 
diagnostic value (L. Brendonck, pers. comm.). 

The type localities are quite different for the 
two species. S. a/r/?m apparently inhabits water- 
holes near sea level that become saline at very 



388 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2. Streptocephalus (Parasireptocephalus) queenslandicus sp. nov. 6 ; A, 3rd right thoracic appendage; B 1, 
detail of 1st anterior setae of first endite; B2, details of 2nd and 3rd anterior setae of 1st endite; C, details of 
anterior setae of 2nd endite; D, details of anterior setae of 3rd endite; details of the anterior setae of 4th endite; 
F, details of antQrior^tae of 5ih endite; G, detail of hook-like setae on median margin of endopodite; H, detailof 
^mftUpliu^o^setae oadhtalmarginof endppodite; I, detail of setae of exopodite; J, detail of hook-like setae on 
octettml margin of exopodite; IQ detail of edge of preepidcpodtte. Abbreviations: end. = endite; endop. = 
endopodhe; exop. — exopodite; pre-^ip. =preepipomte. 



high tides (though there is no suggestion that it 
jives in saline waters), while S. queenslandicus 
lives in fish hateheiy ponds at 590in a.s.l. on fhe 
Athcrton Tablelafld 860km W of Rockhampton- 

While thp di£f^«lices between the femalesof £ 
archert and S. queemtandicus are smali and of 
unknown reliability, the balance of probabilites 



suggest fliey are separate species. Only the re- 
discovery of S. archeri at or near its type locality 
and the recovery of males will provide the basis 
fot amore convincing difibr^tial diagnosis^ 

ECOLOGY, Like most anostracans, S. queem- 
l(mdicus has bem observed to hatch soon (24 



NEW STREPTOCEPHALUS FROM NORTH QUEENSLAND 



389 



hours) after the filling of a 
pond and take 2-3 weeks to 
reach maturity. Hatching can 
occur in any season, with 
growth hardly affected by 
temperature which varies 
annually from 17-28°C. They 
generally persist in the ponds 
for about a month, with males 
dying much earlier than 
females. Only after the ponds 
are drained, dried and refilled 
do they reappear. It is possible 
that dispersal is only by 
resting eggs in mud, as so far 
newly constructed ponds 
using water from ponds 
infested with S. queens- 
landicus have not developed 
populations. 

Shrimps feed on algae and 
do best when phytoplankton is 
abundant soon after hatching. 
They seem incapable of using 
either Volvox or Anabaena 
which often predominate in 
the hatchery ponds. Some- 
times they die out before 
reaching maturity — this 
happened in the October 1997 
hatching and seems to have 
been due to a large reduction 
in the algae in the pond caused 
either by their own feeding or 
by a large population of the 
cladoceran Moina micrura. In 
hatchery ponds S. queens- 
landicus is a serious 
competitor for the more 
desirable M. micrura and 
copepods. It is controlled by 
the use of hydrated lime at 
40-50ppm applied 3-6 days 
after filling of the ponds. 

BIOGEOGRAPHY 







The confirmed presence of Streptocephahis in 
Australia raises interesting biogeographical 
questions. This genus is represented in tropical 
and warni temperate areas of the Neoarctic, 
Palaeartic and especially the African biogeographic 
realms by about 50 species (Banarescu, 1990; 
Beik & Brtek, 1995), and now the Australian 
region has at least two species. Significantly both 
species occur in the tropics. However, while 



FIG. 3. Streptocephalus (Parastreptocephalus) queenslandicus sp. nov.; A, 
ventral view of 6 genital region; B, detail of median process on basal part 
of penis; C, male telson; D, lateral view of 9 abdomen with brood pouch; E, 
9 antenna; F, 9, tetrahedral egg. 



Streptocephahis is the dominant anostracan in 
Afiica and Branchinella uncommon (Banarescu, 
1990), in Australia the position is reversed. 
Geddes (1981, 1983) in his studies of Australian 
anostracans had numerous collections of 
Branchinella but none of Streptocephalus. There 
is no apparent reason for this difterence, but 
perhaps Streptocephalus arrived later in Australia 
than Branchinella and is unable to successfiiUy 
compete with a relatively large endemic fauna. In 
this context Banarescu (1990) believes that 



390 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Branchinella spread from a Gondwana base 
before Streptocephalus. Adding further evidence 
for the later arrival of the Streptocephalidae in 
Australia is the fact that all forms so far found 
belong to the subgenus Parastreptocephalns 
which seems to be a derived form of Strepto- 
cephalus. Or could it be that the tetrahedral eggs 
of this subgenus give it some advantage in 
competition against other species (Brendonck et 
al., 1992), so that only these forms, rather than 
Streptocephalus sensu strictus, have some chance 
in the competition against incumbent dominant 
Branchinella? 

ACKNOWLEDGEMENTS 

We thank Dr Luc Brendonck for his comments 
on the manuscript and Dr M Geddes for access to 
literature. 

LITERATURE CITED 

BAN ARESCU, P. 1 990. Zoogeography of fresh waters, 

Vol. L (Aula-Veriag: Wiesbaden). 
BELK, D. & BRTEK, J. 1995. Checklist of the 

Anostraca. Hydrobiologia 298: 315-353. 
BRENDONCK, L. 1990. Redescription of the fairy 

shrimp Streptocephalus proboscideus 

(Frauenfeld, 1873) (Crustacea: Anostraca). 



Bulletin van het Koninklijk belgisch Institut voor 
Natuurwetenschappen Biologie 59: 49-57. 

BRENDONCK, L., HAMER, M. & THIERY, A. 1992. 
Occurrence of tetrahedral eggs in the 
Streptocephalidae Daday (Branchiopoda: 
Anostraca) with descriptions of a new subgenus, 
Parastreptocephalus, and a new species, Strep- 
tocephalus (Parastreptocephahis) -zuluensis 
Brendonck and Hamer. Journal of Crustacean 
Biology 12(2): 282-297. 

GEDDES, M.C. 198L Revision of Australian species 
of Branchinella (Crustacea: Anostraca. 
Australian Journal of Marine and Freshwater 
Research 32:253-295. 
1983. Biogeography and ecology of Australian 
Anostraca (Crustacea: Branchiopoda). 
Australian Museum Memoirs 18: 155-163. 

LINDER, F. 1 94 1 . Contributions to the morphology and 
the taxonomy of the Branchiopoda Anostraca. 
Zoologiska Bidrag fran Uppsala 20: 101-302. 

SARS, GO. 1 896. Description of two new Phyllopoda 
from North Australia. Archiv for Mathematik og 
Naturvidenskab. Kristiania 18(8): 1-34. 

WILLIAMS, W.D. 1980. Australian Freshwater Life. 
The Invertebrates of Australian Inland Waters. 
(MacMillan: Melbourne). 
1981. The Crustacea of Australian inland waters. Pp 
1101-1138. In, Keast, A. (ed.) Ecological 
Biogeography of Australia. (Junk: The Hague). 



NEW DI$TI0BimON AND HABl l AT DATA FOR THE VULNBRaBLE fYtSO^dDTD, 

QSLAM TORgU4T4. (KLUGE, 19741 

BAR^rEVM. HINBS, DAVIDi H/a<n*«l,]VfieLi©n£ VENZ A^a!? t!ERBSA|fyB5 

Htnes, B.M.. I laiinnh. n.. \ tn/, M. & l.sic. 1 JOflO Oft 50: New distribittilBnpnd habiuiidata 
for the vulncralile n^gopodid, Dclma torquaia (l-vlugc, 1974). Mumtt^i ofthi Queensland 
Mi«eb;»45(2):39U393, Brisbane, ISSN 0O79-S83$, 

The vulncrabk* p\ gopntiid, ndniu nyrijUiUii ( Klugt:, 1 974 ), has been icgaitiLd as contlned to 
scatlLTcd iDculilics in stotuliituslern Quecnslaml Merc wc report oii iiic (ifU iwo instances for 
this species occurring in the southern soclion ofOLiLx-nsland n I'll lual- iw l^cti l-iion-gion. In 
addition, the habitat ai two new localities, Bujiyti Mouniiiin^ Jnd Western Creek ^ire diliiarenL 
from others in which D- tonjmia has befflfi wiGordcd. O iiyrquata, p^^g&p^iSd. 

dldfrihutififi. /hi'''Ui/. Brn^ulow BeU Bioregion 

BofWyM* Wf^^t ^ox 7, PmntLpokaui 4lSJ-i David tiatmah, Outvnsland Furh and 

Tmf^ Syr^ FSKAr Dept of^Wot Jlifst^t^A, 80 Meiers M IndQOfwpUty 4075, 
jmtrd!f(f: U September JQ99 



Delnm forquafa has been regarded :is l( m i I'l ncd 
to scattered localities in southeastern Queensland 
{e.g. Klugc, 1974; Davidson, 1993; Mel arhnKK 
1999; Que^ttSlaj^d Museum records), iu liie area 
between Ulam (23°35'S, ]5(r36T.) near Rock- 
hamplon, the Bunya Mountains (26*'52'S, 
!51"I4'E) and Brisbane's western suburbs 
(27^1 'S, J52'^58'E). Recent systematic veriebraie 
i'auna surveys on public funds in southern 
Qo«S6Q£0^ftll}|I(i^-S- ctal., 1999j baye extended 
the kncnVn range of this species rioffhwest to 
Blackdown Tableland and inland to \\'estern 
Creek. These arc the tirsl n\o records 1br this 
yr-^cics in the soulhcrn section of Queensland's 
Bi'igalow B^t Bioregion ^BBB), 9n area seriously 

Blackdown Tableland National Piiirfe 
(23°48'58"S, 149° 10'56"J5) minimi Sftnd- 
itone plateau some 20dk3rri west t»f wfadstonc 

(23**5rS. 15ri4'E). Here, a specimen of* D 
torcfuaia \\ as captured in a pitt'all trap by 1'. l-,yrc, 
M. Schul/. G. Ford and M Malhiesdn t»n 2M 
November, I9S/7. ll was photographed and 
released (QM slide H NW438 QM). Al Weslem 
C reek Stale Forest (28^04^37*S, l50°53'5rE), 
an area ofrolling sandsttine hills near Millmerran, 
UTi adull D tonjuafu was hand captured by M 
Vcnz on J October 1998. This ^pecuuen was 
lodg^d'^^ thetjueettslantf Mixsetmi '(^66808^^ 

fhi^^dditioaal specimens ol D fon/uata were 
found during these surveys. On Ihc IV June. 
1997, twoZ). torqiiutu were haiid-capturcd by D. 
] fines and A. Fletcher at Qrongab Stale Fut^^t 
125°58'02"S, I52°05M7''E), Via Killch^mt.'Oiie 
was lodged with the Queensland Mu^^iitYi 



(.163 the other released aller tissues were 
taken for genetic studies (University of Quccns^ 
land. Molecular Zool<>gy Laboratory, vial 
number QR1A043). On w S^Wntter 1997. a 
single specimen of D. torqnahi was similarly 
captiued by D. Hannah in Yarraman Stale Forest 
(26-52'5]^S, i5r'50M2"E). Tlie identification 
of this species was continned by Qiieenslimd 
Museum staff It was photographed (QM 
transparency NY 739-761) and released al the 
capture site.A fbttrthZ?. lonfuata specii*ien,'ha«a 
captured by B. Hines and I. Gynther at Bunya 
Mountains Stale Forest ^( 26'^4V'35"*S, 
15P35'46''R) on 30 April 1998. was released 
followin^tifisuc sampling (University of 
<;!i)0eyi^iBTid,-^v^^^ Ifimtm QRFA256). Lastly, a 
.Specimen of /X torqitata was collected by 
Queensland "National Parks and Wildlife slalY 
durinu a fauna sun ev of Btillvard Conservation 
l»aik (':4'\S7\^9^ S, l52''()3'0(rE), near Gin Gin, 
on 2o February 1997. This specimen was alwi 
lod.ued v\ ith ihe Queensland Musciun (J67859). 

The idemillcation of ammals at four of the sit 
localities was confirmed with \ oucher specimens 
registered at the Q^e^nsj^nd Museurn- Idom-^ 
ificiition at the othef tWO localities, (Blackdown 
Tableland and Bunya Mountains), was confirracd 
by muliiple ob^servers ai each site, with two 
obscners m each case having prior, first-hand 
ftxperienpe ip iilentifying this species (IiBl MM 
andMS; BHandlG). In each exattopte^lliiE^atlCin 
and scalaiion of the individuals were cGnsiiSWwl 
with that described in Cogger (1996)- 

*ni«'Vr%«lem and Ela^kiteiwn bicalitieet 
at^ srgni^Gpti|«« (b(iy ore outside the Southeast 



392 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Queensland Bioregion (SEQB), in the southern 
section of the BBB. Tlie only other record for this 
species from outside the SEQB is of a halchhng, 
collected in 1943 at Ulam (Kluge, 1974), in the 
northern section of the BBB. 

HABITAT 

Habitat features of the localities described in 
this paper vary to that documented elsewhere. 
For example, Wilson & Knowies { 1 988) report D. 
torquata as an inhabitant of eucalypl/acacia 
woodland usually associated with rocky outcrops 
on ridges. This description is comparable to the 
Blackdown Tableland, Grongah and Yan'aman 
localities, which support narrow-leafed ironbark 
{Eucalyptus crehra) open forest with grassy 
understoreys on ridges. It is also similar to the site 
where Porter (1998) conducted the only detailed 
investigations into the ecology of A torquata. In 
contrast, the Bunya Mountains and Western 
Creek sites are different from others in which D. 
torquata occurs. The former supports forest red 
gum (E. tereticornis) w oodland, while the latter is 
an area of brigalow (Acacia harpophylla) and 
belah (Casuarina crisfata) forest with a wilga 
(Geijera pcwvijlora) dominated midstorey. Both 
sites are in low lying areas. 

Ditferences were also noted in soil type. At 
Porter's (1998) D. torquata study site soils are 
shallow, stony lithosols on ridges and texture 
contrast soils on slopes. In contrast, those at 
Western Creek are fine-grained, grey, cracking 
clay sediments, while those at the remaining sites 
are dark-brown to black cracking clays of various 
depths. In addition, most of the specimens 
captured by Porter ( 1 998) were sheltering under 
stones. However, his results indicated that rock 
may not be a nccessar>' component of D. torquata 
habitat, as some individuals were captured in 
pitfall traps set away from rock cover. Neither the 
Western Creek nor the Yarraman sites contained 
significant rock components. 

CONSERVATION STATUS 

A review of known localities for D. torquata 
indicates that its current status is justified. D. 
torquata is listed as vulnerable in both The 
Action Plan for Australian Reptiles (Cogger et 
al., 1993) and under the Queensland Nature 
Conservation (Wildlife) Regulation 1994. The 
discover^' of this species at additional sites 
indicates that it is more widespread than has been 
sunnised. How ever, many of the localities in the 
western suburbs of Brisbane are threatened by 
urban development (Czechura Sc Covacevich, 



1985; McDonald etal., 1991). In addition, of the 
six new locahties discussed here, three are in 
communities considered either 'vulnerable'or 
'endangered" (Table 1) (Young, 1999; Young et 
al., 1999). These findings highlight the need for 
fiirther research into the biology and consen'- 
ation requirements of this species. 

ACKNOWLEDGEMENTS 

We thank the field and office stalT involved in 
the fauna component of the Comprehensive 
Regional Assessment program in southern 
Queensland; Jeanette Covacevich and Patrick 
Couper of the Queensland Museum for assist- 
ance with the preparation of this paper; the 
Queensland Dept of Environment and Heritage, 
the Queensland Dept of Natural Resources and 
Environment Australia which frmded the sur\'eys 
on which these specimens were recorded. In 
addition, we thank Queensland National Parks 
and Wildlife Service staff Paul Horton, Adrian 
Kampf and Steve Clark for the Bullyard CP 
specimen and John Hodgon for information 
regarding this record. 

LITERATURE CITED 

COGGER, H.G 1996. Reptiles and Amphibians of 
Australia. 5th Edition. (Reed Books: Sydney). 

COGGER. H., CAMERON. E., S.^DLIER R. & 
EGGLER, P. 1993. The Action Plan for Aust- 
ralian Reptiles. (Australian Nature Conservation 
Agency: Canbeira). 

CZECHURA, G.V. & COVACEVICH, .1. 1985. Poorly 
known rcpliles in Queensland. Pp. 471-476. In 
Grigg, G, SHINE, R. & EHMANN, H. (eds) 
Biology of Australasian frogs and reptiles. 
(Surrey Beatt\ & Sons: Chipping Norton). 

DAVIDSON, C. 1993. Recovery Plan for the Collared 
Legless Lizard (D. lurquaia). ANCA Endangered 
Species Programme. LInpubl. report to 
Queensland Department of Environment and 
Heritage. 

EYRE, T, KRIECJER, G, VENZ, M., HASELER, M., 
HINES, B. & aMVNAH, D. 1999. Systematic 
Vertebrate Fauna Survey Project. Stage I - 
Vertebrate Fauna Survey in the South East 
Queensland Bioregion. Lnpubl. report to the 
Queensland CRA/RFA Steering Committee by 
the Department of Environment and Heritage, 
Brisbane. 

KLUGE, A.G 1974. Taxonomic revision of the lizard 
family Pygopodidae. Miscellaneous Publication 
of the Museum of Zoology, University of 
Michiiiitn No. 147. 

McDonald, k.r.. covacevich, j.a., Ingram, 

G.I. & COUPER, P..I. 1991. The status of frogs 
and reptiles. Pp. 338-345. In Ingram, GJ. & 
Raven, R.J. (eds) An atlas of Queensland's frogs, 



NEWDATA POfe£>£LMi WRQUATA 



393 



TABLE 1. Conservation status and description of Regional l-cosyslems ai D. torqitaia locations. Regional 
Ecosystem numbers and edited descriplions are derived from Young ( 1 999 ) and \'oung ct al. ( 1 999). 



Site 


Regional Eco^fstem 


Extent 
reserved 


Extent 
remaning 


Conser\'ati(Ml 
status 


Blackdovvn 
Tableland NP 


not described 








Buiiyard 


Conmthia ciiyiodom ~ Eucaiyptits acmenoides i Cimermedia, E. fibrosa 
subsp. fibmsct. C tracl^hloia, £. mo/uci^inajloweir slopes),^, cr^bra,E. 
exsi-vta Uill woodland oit complin of l7CfnnBtitxbitbiy'«ra 
scdimenlar} rocks 




Low 


71% 




GfongahSF 


Eucalvptus crebra grassn/ woodland on MeEtosBoic b> ProterozDic igpeous rocks 
(12.117). 


Low 


26% 


Vulnerable 


Bunya Mts 
SF 


Eucalyptus teretlcorms open forest to woodland 0li:C!^QZCUC alluvial 
plains including older floodplain complexes. (12.3,3) 


Ijpw 




^ Vulnerable 


VanGtman SF 


not described 








Wftstero 
Creek SF 


Acacia harpoph> 11a A Castiarina crislata shrubb^f open forest on Calnozoic 
to Proterozoic consolidated, fine grained se(HiBettt.i-o\\ lands. Detrp lexiurc 
contrast soils and cracking clays, often gilgmed- H I -9 5 K 


Low 




Endangered 



xeptUeSi, bird3 ^nd tnamnials. (Queensland 

McFAiaj\l^,Da559.ForestVertefeatB^ 

ibr a C!Qtn;piehepsive Regional Assessment in 
Sputiireast Queensland. S^e IlA: Analysis and 
Reserve OpBon Example. Attachment:- Species 
Summaries.itepcsrt to the Queensland CR.-V RFA 
Steering Committee by the Department of 
Environment, Brisbane. 

NATURE CONSERVATION (WILDLIFE) 
REGLTATION 1994. CJucenslaJid Subordinate 
Legislation 1991 No. 474, Nature Conservation 
Act 

PORTER, R. 1998. Ohserv alions on a large population 
of the vulnerable [\\gopodid. torquuta. 
Memoirs of ihe Queensland Museum. 42(2): 
565-572. 



WILSON, S.K. & KNOWLES, D.G 1988. Australia's 
Reptiles: A Photographic Reference to the Ter- 
restrial R^til^ of Australia, (Collins: Sydqey)^ 

YOUNG, RA.R. Chapter JZ. Southeast 

Queensland; In Sattleii ?A & WlUams^ RJ). 
(eds) The conservation status -of ■QteBeiiMand*s 
bioregional ecosystems. Copserv^oti Technical 
Report. (Queensland Department ofHlvironment 
and Heritage: Brisbane). 

VOLiNG, P.A.R., WILSON, B.A.. McCOSICER, J.C., 
FENSHAM, R..I., MORGAN. G. & TAYLOR, 
RM. 1999. Chapter 11. Brigalovv Belt. In Saltier, 
P.S. & Williams, R.D. (eds) The conservation 
status of Queensland's bioregional eco^\ stems. 
Conservation Technical Report. (Queensland 
Department of Environment and Heritage: 
BrisbaiR^. 



394 



MEMOIRS OF THE QUEENSLAND MUSEUM 



ADDITION TO THE HERPETOFAUNA OF 
QUEENSLAND'S BRJGALQW BIOGEOGRAPHIC 
JtEGlON^J^pirs oftHeQu^en^lftndM^^ 45(2): 394. 
2000:- Th« heipetofauna of the ^igal'ow Biogeographic 
Region has been assessed and documented in some detail 
(Covacevich et al. 1 996). Their work comprised a review and 
synthesis of material held in several museums, collation of 
sight records from numerous auihiiriiies w iiliin Ihe region and 
reports of cxicnsivc llcklwork. One hundred and forh^ five 
species of repliles were recorded a> occurring wilhin die 
Brigalow Biogeographic Region (C n\aee\ich el al. 19^6). 

In December 19^7, 1 located a pre\iousl> unrecorded 
species, Vamniis stwri, in die noilhern portion oftlie region. 
A single specimen was Ibund on the Nehu lo ( ollinsx ille road, 
c.lOkm north ol' (he Ncvvlands tumon' and 50km S of the 
junction witli tlie Mt Coolon - CoUtnsville road at 
148°13'E. The road passes through Homevale and Emu 
Plains. The specimen Wfs located beneath a large fock: pn the 
toad verge (Fig. 1 ). A transparency of the specimen lias been 
lodged with the Queensland Museum images collection (Reg. 
no. C)G37). 

The lowrolling hill area where the specimen was found isa 
mixed eucalypt open woodland of Mountain Coolibah 
{Eucaiyptus atj^dopkila) and f*OT>lar Box (£. popvlma) 
interspersed with open grassland patches of Bluegrass 



{Bothriochloa decipicns, nuJunihtun] scriceum) and 
Kangaroo Gi^s ijh^medu uiLsiruUs), Soils in this ar^ are 
cl^y Ipfuns dorjved from shales.. Some outcropping of these 
shiiles occurs on1^ peadcs dfthe lo\* hills. At the point where 
the specimen was located surface rocks were well embedded. 

Varamis storri is found usualK w here rocks provide cover 
for their shallow burrows (pers. obs.). Abundant surface 
ouicTOpping occurs wilhin 1 0km lo Uie nordio^the site \\ here 
the specimen w as located, thougll this area was not seiirehed. 
U >eenis likeK that rocks on roadside verges m;i\ p[o%ide 
corridors ol suiiahle habitat. V suirri is common in simdar 
habitat approximately 150km west ol'this site, in ^e Desert 
Uplands Biogeographic Region (pers. obs.). 

Aclcndwlcdsekneiiits 

Jplians McCostcer of the Qepartqasnt of ^vuronment, 
Em^ld, identified botanical sainples fh>m Re sile. 

Literature Cited 

COVACEVICH, J.A., COUPER. PJ. & McDONALD, K.R. 1996, 
Replilea of Queensland's Brigftlow Biogeogrfiphic Re^OAt 
Distribution, Status, Conservation. Unpubl. report to 

Australian Nmional Council . 

Steve XIcAlpin, 14 XWLsStreet. A I ice Springs 0870, Australia; 
J 7 May 2000. 




FIG 1 . Varanus storri on the Nebp to ColSnsVitleToad, 



W01 ARD D. BOARE AND ALEX G COtWC 

Hdare, R.D. & Cook. A Ci lOiiO Do .^0: Devonian ?ind Harl> Carhruiirerous I^t^lyplacophora 
Xrqm Western Australia. Memoirs qj the Qm^emlami h'iuaeum 45(2): 395-403. Brisbane. 
ISSN 007^^5. 

Three new taxatdfjp&lyplatophorans are described fi'om tlic C&rboniferous (Toumaisiin) 
Septiijius Lbnesione^d YiseanUtUng CaJcorenUeof the Bonaparte Gulf Basin. Wesleia 
Australia. Mew laxaAtelS^jitectooK cdtiecfus sp. nov,, Compsochironformosus gen. et sp. 

& Scut^eon; 1976 Is described' fmix tw Oemiati Ftasni^n Ss£m I^imestonfi; C^ftnltig 
sft^t^ Western Austral ia, "^l PnlypMc^h^ti^ ptv^^riiipf^ Oftr^^f^apids, $&rtetpertif Gw 
Bas'm. Canning Basin. Hestern Auslralfa. 

Kifhunl D- Hoare. Department of Gcolo^', BmvUng G>'eeH St(tte Umversity, pow/fng 
Grtv-n, Ohio 43403 USA: Alex G. CoQk, Queensland /nfustnim, POBoxJSOQ/SaurhBrhbme 

4Wi Auslnilia: 21 March 2000. 



Plates of Palaeozpic polypiacophorans ha\c 
jlotbeen commonly tqsarNklftetii Australia. The 
first recorded occurrence was Chelodes 
calcenioidcs by Ethcridge (1897), from the 
Upper Silunan of New South Walc^ (NSW). 
Iredale & Hull ( 1^26) described a Pennian taxon 
from NSW, Perniochttan anstt^cdiMUS and 
Farrell (1992) described Chelodes inteijnedius 
Betgenhayn, 1960 and HelmMhochiion sp. nov. 
from the Early Devonian of NSW, The Devonian 
arid Carhoniferoub specimens described herein 
Ironi Wesiem Australia (WA) add subsianiially 
to our knowledge of Palaeozoic Pol^ piacophora. 
in tjiie'c^fb^ta^esfl. 

GEOLOGY And AGE 

The onshore Bonaparte Gulf Basin. WA, 
includes a number of Lowei Carboniferous units 
wliich have yielded a rich and di%'erse fossil 
fauna. Those fatmas suidied include brachropods, 
.(Roberts, 1971i HiQinas, 1971), conodonts 
(Dnic^, 1%9> and oSitea^des (Jones, 1989). 
StUdies of molluscan tjl^iiients of the Car- 
boniferous Kaunas are nowundenvay. The presenl 
W^kis concerned witli polyplaeophoran plates 
njcovered from silicified faunas in the Lining 
Calcarenile on the northwestent shell, and the 
Scptmius Limestone on the eastern shelf of Ihe 
onshore Bonaparte Gulf Basin. For a recent sliirt- 
mary of the stratigraphy and regional geology see 
Mory &. J3eere ( 1 990 ). A dclaiJcd biosti'^igrapliic 
summary vvJisproVi*!^^ 

The Septimus Limestone crops-out in the SE of 
the Bonaparte Basin and has been assigned a 



(Uisfrali.s /.one to lower Spirifer spirt/is Zonc^ 
prmiarily based on bracliiopod faunas (Rob«5t1Si 
1971). Chitons described from the Septimus 
Limestone are from the upper pcul of the unit cmd 
are hcnec regarded as Tn-, in age. The L'lting 
Calcarenile is a laterally restricted unit which 
crops out in the N W of the onshore basin and has 
been assigned a Visean (.Vab, J 3gev Punctospirifcr 
paiiclplicahis Zone. 

In addition to the Carboniferous ta\a. a new 
Late Devonian occurrence is described. The single 
plate was recovered from the Late Devonian 
(Frasnian) Pillara Limestone, in the Hull Range, 
Canning Basm, WA. All material is held in the 
Queenslatid MuseuTQ P^l^eofitQlogy a)UectiQn$ 
(QMF). 

List of Localities, 0ML1^5 Utting Cateateftute. 

Uttine Ciap. Ningbine Station, Bonaparte Gulf 
BasirC WA. 14'\S8.17''S- ]28^15,S2"E, Coll A. 
Cook P. Jell, May 1996: A Cook, T-Smiih, July 
1998. QMLi096 SqDiimus Limestone, low spur 
on NW side of Mt Septimus. Ivanhoe Station. 
Bonaparte Gulf Basin. WA, 15M2.5^S, 
1 28^'59.22T.. Coll. A. Cook, R .Fell. May 1 996; A. 
rook. T Smith, July 1998. OML1160. Sadler 
Limestone. SW side of Paddy's Valley, SSL 
Wade Knoll Camnri^i Basin, WA, 18W07" S. 
125^59 3 l"E. CqIL A. Cook. T. Smith, This is 
near Stop X rtf P«ay^098l). 

the reconsmtctiotig (Pig. D ltltiStratc p»:tie 

rehitionships in the three Carboniferous ta.xa 
described herein. Head plates arc unknown ft>r 
Gi^biticMt6nsdliec1us^,VX^ 



396 



MEMOIRS OF THE QUEENSLAND MUSEUM 






FIG. 1 . Reconstructions. A, B, Gn^phochilon collectns sp. nov., dorsal and right lateral views, approx. X 1 .0. C, 
D, Compsochiton formosus gen. et sp. nov., dorsal and right lateral views, approx. X 1 .0. E, F, Harpidochiton 
aiictus, gen. et sp. nov., dorsal and right lateral views, approx. x 1 .0. 



auctus gen. et sp. nov., but the number of other 
plates present for each taxon allows some basis in 
forming arrangements as presented. No inform- 
ation as to size of girdle or presence of girdle 
spines or plates is present. 

SYSTEMATIC PALAEONTOLOGY 



MATERIAL. HOLOTYPE: QMF51013, from QML1096, 
Septimus Limestone, Toumaisian, Bonaparte Gulf Basin. 
PARATYPES: QMF5 101 4-5 1022 from QML1096. 

DIAGNOSIS. Tail plate narrowly elongate; 
intermediate plate elongate, wider posteriorly 
than anteriorly, strongly arched. 



POLYPLACOPHORA de Blainville, 1816 
LEPIDOPLEURIDAE Thiele, 1910 
GRYPHOCHITONIDAE Pilsbry, 1900 
Gryphochiton Gray, 1 847 

Gryphochiton collectus sp. nov. 
(FigslA,B, 2) 

ETYMOLOGY. Latin, collectns. narrowed, contracted. 



DESCRIPTION. Plates of moderate size (Table 
1). Tail plate elongate, narrow, strongly arched. 
Jugal area prominently set off from lateropleural 
areas, mucro small, j ust anterior to posterior mar- 
gin. Posterior margin slightly arched; anterior 
margin straight. Sutural laminae relatively wide, 
long. Surface smooth with prominent comarginai 
growth ridges posteriorly and laterally. Angle of 
jugal area 20°. 



DEVONIAN AND EARLY CARBONIFEROUS POLYPLACOPHORA 



397 




FIG. 2. Gryphochiton collectiis sp. nov. A-C, Holotype QMF51013, tail plate, dorsal, ventral and right lateral 
views; D, Paratype QMF5 1 020, tall plate, dorsal view; E,F, Paratype QMF5 1015, intermediate plate, dorsal and 
ventral views; G, Paratype QM F5 1018, intermediate plate, dorsal view; H-J, Paratype QMFS 1014, intermediate 
plate, dorsal, right lateral and posterior views; K,L, Paratype, QIVIF5 1016, intermediate plate, dorsal and ventral 
views; M, Paratype QMF51018, intermediate plate, dorsal and right lateral views; P,Q, Paratype QMF51019, 
intermediate plate, dorsal and anterior views. All x 2.5 



398 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Measurements for Gryphochiton collectus 
sp. nov. 



Specimen 


Length 
(mm) 


Width 
(mm) 


Height 
(mm) 


Plate 


QMF510I3* 


LI. 5 


7.8 


2.4 


T 


QMF51014 


16.0 


12.5 


5.9 




QMF510I5 


12.9 


9.2 


3.9 




OMF51016 


14.9 


11.6 


5.4 




QMF51017 


18.5 


11.8 


5.5 




QMF510I8 


13.7 


10.2 


4.7 




QMF51019 


10.6 


9.2 


4.3 





Intermediate plates longer than wide, tapering 
anteriorly, strongly arched. Lateral margins 
gently convex, anterior margin straight to slightly 
convex, posterior margin slightly mucronate. Jugal 
area distinct from lateropleural areas. Apical area 
large, extending as broad band along posterior 
margin. Surface smooth with prominent co- 
marginal growth ridges on lateropleural areas. 
Apical angle 94° to 112", angle otjugal area 18" to 
23". Headplate unknowTi. 

REMARKS. Gryphochiton collectus sp. nov., 
although smaller, is most similar to G. nei-viccmus 
(de Ryckholt, 1845), the type species of 
Gryphochiton Gray, 1847b. Comparisons were 
made with three tail plates and one intermediate 
plate of G. nerviccmus in the collections of the 
Museum of Comparative Zoology, Harvard 
University. The major difference lies in the much 
larger sulural laminae, larger apical areas, and 
less strongly arched posterior margin of the tail 
plate in G. collectus. 

Compsochiton gen. nov. 

TYPE SPECIES. Compsochiton formosus sp. nov. 

ETYMOLOGY. Greek, kompsos, elegant, pretty; chiton, 
tunic. 

DIAGNOSIS. Tail plate with broadly angular 
jugum, posterior margin flatly convex and curved 
dorsally; intermediate plate mucronate; sutural 
laminae large. 



DISTRIBUTION. Carboniferous (Toumaisian), 
Septimus Limestone, Bonaparte Gulf Basin, WA. 

REMARKS. Compsochiton differs from Grypho- 
chiton Gray, 1 847a by having differentiation of 
jugal and lateropleural areas on the head plate, a 
small falsebeak on the tail plate, and lack of the 
subquadrangular to subrectangular shaped inter- 
mediate plates of the latter genus. Euleptochiton 
Hoare and Mapes, 1985, has a much wider than 
long tail plate with a narrower jugal area, and 
mucro located anterior to midlength. 

Compsochiton formosus sp. nov. 
(Figs 1 CD, 3) 

ETYMOLOGY. LdJcin^ formosm, beautifully formed. 

MATERIAL. HOLOTYPE: QMF50I23, from 
QML1096, Septimus Limestone, Tournaisian, Bonaparte 
Gulf Basin. PAR\TYPES: OMF50 124-501 36, from 
QML1096. 

DIAGNOSIS. As for the genus. 

DESCRIPTION. Plates of moderate size (Table 
2). Tail plate strongly arched with broadly 
angular jugal area, mucro low and posterior to 
midlength. Posterior margin flatly convex, lateral 
margins diverging anteriorly, anterior margin 
with small false beak. Jugal area slightly set oft^ 
from lateropleural areas, the latter marked by 
distinct comarginal growth ridges. Sutural 
laminae wide, long, extending to anterolateral 
comers. Surface ornamented with fine, closely 
spaced pustules. Angle of jugal area 70°. 

Intermediate plates wider than long, strongly 
arched, with broadly angular jugal area, slightly 
to distinctly set off from lateropleural areas. 
Posterior margin mucronate, lateral margins 
convex, anterior margin ranging from slightly 
convex to slightly concave. Apical area large 
medially, extending as narrowing band to 
posterolateral comers. Surface with fine, closely 
spaced pustules and distinct comarginal growth 
ridaes on lateropleural areas. Apical anale 93" to 
106^ angle of jugal area 48° to 60^ 

Head plate strongly arched with broadly angular 
jugal area distinct from lateropleural areas. 



FIG. 3. Compsochiton formosus gen. et sp. nov. A-E, Holotype QMF51023, tail plate, dorsal, ventral, left lateral, 
anterior and oblique ventral views; F,G, Paratype QMF5 1 024, tail plate, dorsal and ventral views; H, Parat>'pe 
QMF51026, tail plate, dorsal view; I-L, Paratope QMF51028, intermediate plate, dorsal, ventral, right lateral 
and posterior views; M-0, Parat>'pe QMF51027, intermediate plate, dorsal, ventral and anterior views; P,Q, 
Paratype QMF5 1 029, intermediate plate, dorsal and right lateral views; R, Parataype QMF5 1 030, intemiediate 
plate, dorsal view; S,T, Paratype QMF51031, intermediate plate, dorsal and left lateral views; LI,V, Parat>'pe 
QMF5 1 033, intermediate plate, dorsal and ventral views; W,X, Paratype QMF5 1 034, head plate, dorsal and left 
lateral views; Y-AA, Paratype QMF5 1035, head plate, dorsal, ventral and posterior views. All X 2.5 exceptE x 4. 



DEVONIAN AND EARLY CARBONIFEROUS POLYPLACOPIIORA 



401 



TABLE 2. Measurements for Compsochifon formosus 
gen. et sp. nov. 



specimen 


Length 
(mm) 


Width 
(mm) 


Height 
(mm) 


Plate 


QMF50123* 


9.3 


9.7 


5.2 


T 


OMF50124 


9.4 


9.4 


3.8 


T 


OMF50125 


8.7 


9.0 


4.2 


T 


QMF50I26 


8.4 


8.7 


4.7 


T 


gMF50127 


9.8 


12.6 


5.5 


1 


QMF50128 


10.3 


14.0 


6.7 


I 


QMF50129 


9.5 


11.1 


5.1 


I 


QMF50130 


9.4 


11.4 


5.6 


1 


QMF50131 


10.0 


9.9 


5.2 


I 


QMF50132 


9.5 


12.0 


5.8 


I 


OMF50133 


8.8 


11.2 


5.3 


I 


QMF50134 


6.3 


9.2 


5.1 


H 



Anterior and lateral margins broadly curved, 
posterior margin nearly straight. Apical area 
large, extending along posterior margin to 
posterolateral corners. Surface with fine pustules 
and distinct comarginal growth ridges in latero- 
pleural areas. Apical angle 106**, angle of jugal 
area 60". 

REMARKS. At first appearance C formosus 
looks similar to Gryphochiton parvus (Stevens, 
1 858), although the plates are three times the size 
of the latter species. Other than size, the distinct, 
broadly angular, jugal area, large sutural laminae, 
false beak on the tail plate, and the finely 
pustulose ornamentation on C. formosus clearly 
differentiate the two taxa. 

INJURY. The holotype of Compsochiton formosus 
gen. et sp. nov., a tail plate, has an injury in the 
posterolateral margin (Fig. 3B, C, E). A notch, 
approximately 0.75mm wide and 1.5mm high, 
extends through the shell material. It is evident 
that the chiton was alive at the time of the injury 
by the presence of new shell material deposited 
on the ventral surface to block off the opening. 
No indication is present as to the cause of the 
injury although the bulging inward shape of the 
repair made by the individual would indicate that 
either the mantle at the site had been injured or 
that the predator was present within the notch in 



the original shell layers. This is the first example 
of plate injur>' and repair of a fossil poly- 
placophoran that we are aware of. 

LEPIDOPLEURIDAE? Pilsbry, 1892 

HarpidochitoD gen. nov. 

TYPE SPECIES. Harpidochiton auctus sp. nov. 

ETYMOLOGY. Greek, harpidos, shoe or sandal; chiton, 
tunic. 

DIAGNOSIS. Tail plate subtriangular, longitud- 
inally concave, mucro tenninal; intermediate 
plates subquadrangular, strongly arched, apical 
area large. 

DISTRIBUTION. Carboniferous (Toumaisian), 
Septimus Limestone, Bonaparte Gulf Basin, 
Western Australia; (Visean) Utting Calcarenite, 
Bonaparte Gulf Station, WA. 

REMARKS. Harpidochiton differs from genera 
of the family Acutichitonidae Uoare, Mapes & 
Atwater, 1983, also with subtriangularly shaped 
tail plates, by lacking a hypotyche on the ventral 
surface. Systenochitoti Hoare (in press), Irom the 
Mississippian (Osagean) of Iowa lacks the 
longitudinal concavity in the subtriangular tail 
plates, has a subcentral mucro, and has much 
smaller sutural laminae than is present in 
Harpidochiton. 

The characters of Harpidochiton do not conform 
well with those of the other lepidopleurids. nor 
with other described families. It is assigned here 
temporarily pending additional material on 
which to base a new family. 

Harpidochiton auctus sp. nov. 
(Figs IE, F, 4) 

MATERIAL. HOLOTYPE: OMF50137, from 
QML 1 096, Septimus Limestone, Toumaisian, Bonaparte 
Gulf Basin. PARATYPES: QMF5()138-50144, from 
QML1096, QMF40826-40829, from QML1095, Utting 
Calcarenite, Visean, Bonaparte Gulf Basin. 

DIAGNOSIS. As for the genus. 

DESCRIPTION. Plates of moderate size (Table 
3). Tail plate subtriangular. Lateral margins con- 
vex, anterior margin straight to weakly convex. 



FIG. 4. Harpidochiton auctus gen. et sp. nov. A-C, Hoiot>pe QMF5 1 037, tail plate, dorsal, ventral and left lateral 
views; D-F, Parat>pe OMF51038, tail plate, dorsal, anterior and ventral views; G, Paralype OMF51039, tail 
plate, dorsal view; H-J, Paratype QMF5 1 040, intermediate plate, dorsal, right lateral and posterior views; K-M, 
Paratype QMF5104I, intermediate plate, dorsal, ventral and anterior views; N, Paratype QMF51043, 
intermediate plate, dorsal view; 0,P, Paratype QMF51042, intermediate plate, dorsal and left lateral views; 
Q-T, Paratype QMF40826, intermediate plate, dorsal, ventral, right lateral and posterior views; U,V, Paratype 
QMF48027, intennediate plate, right lateral and dorsal views. All x 2.5. 



402 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 3. Measurements for Harpidochiton auctus 
gen. et sp. nov. 



specimen 


Length 
(mm) 


Width 
(mm) 


Height 
(mm) 


Plate 


QMF50137* 


10.9 


ll.O 


5.9 


T 


QMF50138 


13.8 


13.2 


7.9 


T 


QMF50139 


10.8 


12.1 


7.0 


T 


QMF50140 


13.7 


13.1 


7.0 


I 


QMF50141 


11.6 


11.2 


6.0 


I 


QMF50142 


10.7 


12.3 


5.6 


I 


QMF50143 


11.1 


12.9 


6.5 


I 



Jugal area set off from lateropleural areas by 
pronounced grooves; mucro terminal. Lateral 
profile longitudinally concave. Sutural laminae 
large. Surface smooth with strongly developed 
comarginal growth ridges on lateropleural areas. 
Angle of jugal area 33°. 

Intennediate plates subquadrangular, strongly 
arched, sharply curved transversely in jugal area. 
Lateral margins gently convex, anterior margin 
with wide, shallow jugal sinus, posterior margin 
straight to slightly mucronate. Jugal area promi- 
nent, set off by grooves. Sutural laminae large, 
extending to anterolateral comers. Apical area 
large, extending to posterolateral comers, otlen 
marked by comarginal growth ridges. Surface 
smooth with prominent growth ridges on latero- 
pleural areas. Apical angle 93° to 103°, angle of 
jugal area 35° to 39°. Head plate unknown. 

REMARKS. The concavity of the tail plate, the 
smooth surface, and the coarse, comarginal 
growth ridges restricted to the lateropleural areas 
are diagnostic of H. auctus. 

ACUTICHITONIDAE Hoare, Mapes 

& Atwater, 1983 
Arcochiton Hoare & Sturgeon, 1976 

Arcochiton sp. 

(Fig. 5 ) 

MATERIAL. One tailplate QMF39839 from QML1160. 

DESCRIPTION. Small (3.2mm long, 3.3mm wide, 
2.4mm high) tail plate, subtriangular in dorsal 
and lateral views. Jugal area narrow, slightly set 
off from steep lateral slopes, mucro terminal 
posteriorly. Anterior margin with deep jugal sinus, 
large, well-developed hypotyche on ventral 
surface. Surface granulose, augmented by silica 
replacement, especially near lateral margins. 
Evidence of narrow sutural laminae on ventral 
surface paralleling margins of jugum, their pro- 
jections not preserved. 




FIG. 5. Arcochiton sp. A-D, Hypotype QMF39839, tail 
plate, dorsal, ventral, left lateral and anterior views. 
All X 10. 



REMARKS. The presence of this Australian 
specimen extends the known range of the family 
Acutichitonidae from the Permsylvanian to the 
Upper Devonian and specimens from Oregon 
further extend that range into the Permian 
(Hanger et al., 2000). The coarse preservation 
partially masks the characters of the plate. The 
shapes and curvature of the plate and hypotyche 
are distinctive of Arcochiton. More and better 
preserved material would allow the establish- 
ment of a new species. 



DEVONIAN AND EARLY CARBONIFEROUS POLYPLACOPHORA 



403 



LITERATURE CITED 

BLAINVILLE, H.M.D. De 1816. Prodrome d'une 
nouvelle distribution systematique du regne 
animal. Bulletin Sciences Societe Philomathique 
de Paris Pp. 105-124. 

BERGENHAGEN, J.R.M. 1960. Cambrian and 
Ordovician loricates from North America. Journal 
of Paleontology 34:168-178. 

DRUCE, E.G. 1969. Devonian and Carboniferous 
conodonts from the Bonaparte Gulf Basin, 
northern Australia. Bureau of Mineral Resources. 
Geology and Geophysics Bulletin 98: 1-243. 

ETHERIDGE, R. Jr 1897. On the occurrence of the 
genus Chelodes Davidson and King, in the Upper 
Silurian of New South Wales. Records of the 
Geological Survey of New South Wales 5(2): 
67-70. 

FARRELL, J.R. 1992. The Garra Formation (Early 
Devonian: late Lochkovian) between Cumnock 
and Larras Lee, New South Wales, Australia: 
stratigraphic and structural setting, faunas and 
community sequence. Palaeontographica Abt A 
222: 1-32. 

GRAY, J.E. 1847a. On the genera of the Chitonidae. 
Proceedings of the Zoological Society of London 
179: 63-70. 

1 847b. A list of the recent Mollusca, their synonyms 

and types. Proceedings of the Zoological Society 

of London 179: 168-169. 
HANGER, R.A., HOARE, R.D. & STRONG, E.E. 

2000. Permian Polyplacophora, Rostroconchia, 

and Problematica tYom Oregon. Journal of 

Paleontology 74(2): 192-198. 
HOARE, R.D. (in press). Early Mississippian 

Polyplacophora (Mollusca) from Iowa. Journal of 

Paleontology. 

HOARE, R.D. & MAPES, R.H. 1985. New Missis- 
sippian and Pennsylvanian Polyplacophora 
(Mollusca) from North America. Journal of 
Paleontology 59(4): 875-881. 

HOARE, R.D., MAPES, R.H. & ATWATER, D.E. 
1983. Pennsylvanian Polyplacophora (Mollusca) 
from Oklahoma and Texas. Journal of 
Paleontology 57(5): 992-1000. 

HOARE. R.D. & STURGEON, M.T \ 97 6. Arcochiton 
raymondi n. gen., n. sp. of polyplacophoran from 
the Pennsylvanian of Ohio. Journal of Paleont- 
ology 50(5): 841-845. 

IREDALE, T & BASSET HULL, A.F. 1926. A mono- 
graph of the Australian Loricates (Phylum 



Mollusca-Order Loricata). Australian Zoology 
4(5): 324-328. 

JONES, P.J. 1989. Lower Carboniferous Ostracoda 
(Beyrichicopida and Kirkbyocopa) from the 
Bonaparte Basin, northwestern Australia. Bureau 
of Mineral Resources, Geology and Geophysics 
Bulletin 228: 1-97. 
1 993. Early Carboniferous Cladocopina (Ostracoda) 
from the Bonaparte Basin, NW Australia. 
Memoirs of the Association of Australian 
Palaeontologists 18: 143-152. 

MOREY, A.J. & BEERE, GM. 1990. Geology of the 
onshore Bonaparte and Ord Basins in Western 
Australia. Geological Survey of Western 
Australia Bulletin 134: 1-184. 

PILSBRY, H.A. 1892. Monograph of the Poly- 
placophora. Manual of Conchology, Academy of 
Natural Sciences, Philadelphia 14. 

PILSBRY, H.A. 1900. Amphineura. Pp. 433-436. In 
Zittel, K.A. (ed.) Textbook of paleontology, 
Eastman translation. (Macmillan & Co: New 
York). 

PLAYFORD, P.E. 1981. Devonian reef complexes of 
the Canning Basin Western Australia. Field 
Excursion Guidebook, Fifth Australian Geol- 
ogical Convention (Geological Society of 
Australia: Perth). 

ROBERTS, J.R. 1971. Devonian and Carboniferous 
brachiopods from the Bonaparte Gulf Basin, 
northwestern Australia. Bureau of Mineral 
Resources, Geology and Geophysics Bulletin 
122: 1-319(2 vols). 
1985 (ed. Australia). Pp. 9-145. In Diaz, CM. (ed.) 
The Carboniferous of the World 2: Australia, 
Indian subcontinent, South Africa, South 
America and North Africa. lUGS Publication 20. 
(Institute Geologico y Minero de Espana: 
Madrid). 

RYCKHOLT, R DE. 1845. Resume geologique sur le 
genre Chiton Lin. Bulletin de FAcademie Royale 
des Sciences et des Belle-Lettres de Bruxelles 12: 
36-42. 

STEVENS, R.P. 1858. Descriptions of new Carbon- 
iferous fossils from the Appalachian, Illinois, and 
Michigan coalfields. American Journal of Science 
and Arts, Series 2, 25: 258-265. 

THIELE, J. 1909-1910. Revision des Systems der 
Chitonen. Zoologica 22(56): 1-132. 

THOMAS, GA. 1971 . Carboniferous and Early Permian 
brachiopods from western and northern Australia. 
Bureau of Mineral Resouces, Geology and 
Geophysics Bulletin 56: 1-277. 



MfeSOZbie FRISHW^BR AND ESTtlARlNE fiJVALVISfROM AUSTRALIA 



scon A HOfKNDLI, 
ItoerttHUil, S.A. 2O00 06 30: Mcsozoio frcsiiwatcr iuid ostuarine bivulvcs fruii* AuMtraha. 

Auslralian- freshwatet and L-sUiaiiiiL; bivalves artr iltiiict ibcd Irom ihc Hyrridfle;. 
Cilauconomidae ;md Sphacriidae. The following taxa are no-U' kmiwn lioiii the Mcsozolo^jf 
Avstvali.r (Voin the Triassic, ^iyriidac, coitiprising /VoZ/jr/ti trm/.v^.v, Mc^^a/tA'irgns 
nov.- X'kxa/avirgas ckiUmdi, Me^alnvirgus jaemchi and Mi'SohyrkicHa ipsviaeffsis, 
CJIauconomidac, UHioih'Uu wiarumatii^nsis^ ProtovirgitH dwistani and Pmtovif^us 
b/vokvuhfwx nOv.;fwro ijlt: Jurassic;, 1 lysl]j^aQiNyndeUa (PratnljyridelkO sp. Prohyria 
skepUcBi «p. i!t>v., ?SpfiBeritdac, Pt'otosj^^fifm icilbragatemis gen. ct sp. nov. acid 
^Tqtasgkaerium glanae Jtp. nov.; ami, itom the Creiacepu*, Hyriidac, K'fe^aiovjr^m 
Mmotjtnm^ llyridalta macmichaeli, flyrideUn (J'rotBhyridelfa) gouridiwtndiinAi.y. 
ff^n'Idcllc^M^hUecll/fsensfs.AlylharicncHfnc!! 

gbn.elsp. nov. It Is apparent iluit o\ cr t!ic cv)iirse of freshwater bivalve ovohiHon in AusTralia 
thisrc has been a distinct f;iiin;il dichoioniv between large and small ta.\a. I he lurgc loims ;irc 
exclusively hyriids and donuniue lacusiriiie, rj\^ciiiie and iagoonal en\ iionnicnis and the 
small Conns, L-iiher ^laueonomids, splueriids and'or cori:>iruliiis domin.iiie blackish and 
cstuai tiie cm (UMiinem:-. Tliis diclu'toriiy is alsci prcseni in ihe modern csitiarine and 
lrc:>h\vaier !:i\a. The presence ol \:laueoiiamids during the Tri;issic ix ilie i'w-^l and earliest 
record, pnu iding o^^^^d i'\ idcnre (or e'.tuarine deposilional em iroianeni ol lhe VViananiulla 
Shales. Ssdiies Basin. I hr-i is al>o the earliesl record of sphaeriid^ in Australia. Lntle is 
known of the evolntionary relaliiniships witiitn these grtnin^ due lo tIu;ii ^onsri v.iti\'c 
morpholog}-. 3 Mcgulovirgui- ,y:ai. no v. Puluvuhyrkldia ^^udthclpi yyt:. . iu>\ /-V. ■;>} ria 
skiipticsi sp. nav.,ProwsphficritiU7i taJf^r^isut'enais gen, ctsp. nm Pr<?fci^'piw€*'n^tfJ gotnoe sp. 

UoCkrwH. S.A. (email: scofjhia^qm.ciIdgMOU). Jm-'ericbnUa Pulaconioh^^ QutftSnSi}and\ 

M^^m^ m mJ^QQ SmtkMsbm il^^JjcMstrgiio: 29 October 1 m. 



Australian fossil freshwater and esiiiarine mol- 
luscs are poorly know a. w ith i'cw ijublicaiions on 
ihe higher taxonomy ot preseiii lins and extinct 
taxa. A review is offered here toi* the Mesozoic 
ffeshvvatei* Wvalves comprising the fimiilics 
! lyriidae, Glauconomidae aiid Sphacriidae, with 
ledescription ofprcviously described species and 
description oi new tornis. Detailed illustrations 
and measurements are provided so that further 
collection does not ronfb^p Issues of variation 
and taxonomic position. 

Etbertdgo (1§88^ i§92) descdM jevetal 
foniis mm. Atisttgifia assigning most to Unio, 

apart from ihc smaller formsto wiiich he assigned 
Ufiinneila Newton (1915) described ne\N 
Cretaceous unionoids from Lightning Ridge and 
WTiile Cliffs, abo allocating these to Umo.^ Al 
this point the Unionoida compris^' th^^Unfen*- 
idaejMargretifidae and Mulelidae. 

Latcn McMichacI {IS)56) revised laheridge 
.Tr's and Newlon's material, picsenting detailed 
descriptions of fossil nonmarinc l.axa from 
Australasia in an effort to alleviate some of the 



cctnsen ative gfoups. McMichael proposed that 
these fossil uniotioids were from a Muteluiae 
lineage, and included the modem faima within 
this fitqily itva reg^nal revision (McMichael & 
HiscDck, 1^8), Most of MfcWfi^ael'fe ibftaSl 
descriptions were paraphrased froiji^uipf idgc 
.It's and Nt-wton's ciirlicr work smfl li^e 

McMichael (l'-)?6) did. howe\er. erect three 
mostly Mesozoic genera of mutelids: Pmhyrta^ 
Mesohyrfdella and Proiovirgus, neally 

tjfxa. Lwdwaok ( 1 951) describe new taxa from 

the Triassic Leigh Creek Formation, South 
Ausiralia,, proposing that the genus Ihiiu wun 
\alid fur some Australian unionoids and thatthiS' 
confirmed the presence of U niomdac in Australia* 
1 provide evidence to the contraiy. Ludbrookalso 
assigned one species to McMichael *s PrtAovfrgui^^. 
which is now placed within Alaihyria. 

Parodiz & Bontietto ( 196?) provided ev idence 
fora separate endemic ( iondwarum lineage on ilie 
presence and moiphology of the glochidial latval 



406 



MEMOIRS OF THE QUEENSLAND MUSEUM 



radiation was distinct enough to allow familial 
uniquity and sevLM'al taxa in South America were 
also assignable to this new group, Hyriidae, 
instead of the dominant Mutelidae. These con- 
clusions suggest that the unionoids from Australia 
are mostly Gondwanan in origin vsith recent, 
presumably Pleistocene - Holocene, invasions 
from southeast Asia. This view is supported by 
the present study. 

Since McMichaels' review, more material has 
come to light that has shed new and interesting 
information on the taxonomic validity and 
position of the fossil taxa. Shell characteristics 
thai McMichael & Hiscock (1958) put forward 
for the subfamilies within the Hyriidae have also 
assisted herein in allocating the fossil taxa to 
these subfamilies. 

Smaller Triassic bivalves (Unionella) describ- 
ed by Etheridge Jr (1888) and reviewed by 
McMichael (1956) from the Sydney Basin and 
Bowral posed problems for McMichael with his 
only suggestion that they were Triassic hold- 
overs of the Permo-Carboni fcrous family 
Anthracosiidae. This family has a conspicuous 
presence in North America (Rogers, 1965), 
United Kingdom (Weir, 1960) and the former 
Soviet Union (Haas, 1969) diu-ing the Carbonif- 
erous and Pemiian. 1 place these small bivalves 
with the modem family Glauconomidae, but do 
not aile out distant phylogenetic links to the 
anthracosiids. There is a strong possibility that 
convergence has been acting on both these 
groups, indeed all freshwater and estuarine 
bivalves reported here. However, enough 
characteristics have been found that ally these 
small Triassic bivalves to the Glauconomidae. 

Most recently, Hocknull (1994) described a 
new taxon from the Late Triassic of southeast 
Queensland allocating it to Protovirgus, here 
revised to Megalovirgtis gen. nov. Hocknull 
(1997) described new taxa from the Cretaceous 
of Queensland placing all taxa within previously 
recognised genera. The understanding that there 
is considerable confusion over the taxonomy of 
the unionoids and the presence of unrecorded 
families of small freshwater faunas prompted the 
present review. 

New material from the Talbragar Fossil Fish 
Beds, New South Wales, indicate the possible 
presence of Sphaeriidae in Australia. Hampered 
by the lack of well-preser\'ed specimens the 
allocation of the small bivalves to Sphaeriidae is 
tentative. 



As there is an apparent dichotomy between 
bivalve faunas, depositional environment, their 
sizes and phylogenetic history, it remains to be 
seen whether the assignment of different sized 
faunas should lie at the family level. Such 
possibilities are discussed later. Fossils are given 
the prellx SAMF (South Australian Museum), 
QMF (Queensland Museum), UQF (University 
of Queensland) and AMF (Australian Musemn) 
and are deposited in their respective institutions. 

AGE AND STRATIGRAPHY 

Ten major sites, containing nonmarine bivalves, 
are recognised from (Queensland (Qld), New 
South Wales (NSW) and South Australia (SA) 
(Fig. 1 ). The hyriids arc restricted in the Triassic 
to the Ipswich Coal Measures, Qld and the Leigh 
Creek and Springtield Basins, SA. During the 
Jurassic they occur sporadically in the Waloon 
Coal Measures, Warwick, SE Qld and the K.oon- 
warra Fossil Bed. South Gippsland, Victoria. In 
the Cretaceous the family is abundant throughout 
the entire Eromanga Basin, especially the 
Coreena, Griman Creek and Winton Fomations 
ofnorthemNSWand Qld. 

Small freshwater and estuarine bivalve fossil 
faunas (glauconomids and sphaeriids) are restrict- 
ed in the Triassic to the Wianamatta Shales, 
Sydney Basin, and in the Jurassic to the outlying 
Talbragar Fossil Fish Bed derived from the Pur- 
lawaugh Formation, No small nonmarine bivalves 
are as yet known from the Australian Cretaceous. 

There are three major stratigraphic units, in 
which Triassic freshwater bivalves are preserved. 
In SE Qld, the units containing hyriids are the 
Blackstone and Tingalpa Formations, both 
considered Late Triassic (Camian) in age by de 
Jersey (1975) and de Jersey & Hamilton (1965), 
respectively. Both formations consist of fine 
grained to coarse-grained sandstones and 
siltstones/shales, with most of the bivalves being 
derived from the fmer carbonaceous siltstones 
and shales. Preservation is good, with most 
individuals having both valves intact and fine 
ornamentation preserved. The valves are 
generally external moulds with little internal 
morphology preserved. The cavities have been 
replaced with ferrous sediment. The presei*vation 
of both valves suggests an in situ taphonomy. 

Play ford & Dettman (1965) ascribe the Leigh 
Creek Formation, SA, a Late Triassic age on the 
basis of similar palynological floras to those in 
Ipswich. Whilst other authors ascribe ages 
between Late Triassic (Rhaetic ) to Early Jurassic 



MESOZOIC BIVALVES FROM AUSTR.'\LIA 



407 




FIG. 1. Map illustrating Australian localities for 
Mesozoic freshwater and estuarine bivalve fossils. 



(Liassic) on the basis offish faunas (Wade, 1 953), 
plant floras (Chapman & Cookson, 1926) and 
unionoids such as Prohyria eyrensis (Ludbrook, 
1961). The Springfield Coal Basin, SA, is also 
attributable to the Late Triassic from diagnosis of 
equivalent sediments to the Leigh Creek 
Formation (Johnson, 1960) and, again, the 
presence of the distinctive Dicroidiuni and 
Thhmfeldia floras with the presence of the hyriid; 
Prohvria evrensis (= Unio springfieldensis 
Ludbrook, 1961). 

Like the Blackstone Formation, the bivalves 
are preser\^ed as moulds with both valves intact. 
The deposit is interpreted as in situ. 

The Brookvale Quarry and Bowral Quarry 
expose sections through the Wianamatta Shales, 
Sydney Basin, NSW. Small valved glaucono- 
mids occur at all sites and are attributable to the 
Late Triassic, being derived from the Hawkes- 
bury Sandstone and Wianamatta Group 
(Lovering & EIroy, 1969). The shells are pre- 
served as whole specimens or as singular valves. 
Mass accumulations suggest fast burial and 
aggregation of these taxa in the bottom sediment. 
Little is known of the aggregation behaviour/ 
taphonomy of modem glauconomids, therefore. 



no interpretation of the depositional environment 
is given here. 

There is one occurrence of small freshwater 
bivalves in the Jurassic of eastern Australia from 
the Talbragar Fossil Fish Beds, derived from the 
Purlawaugh Formation. The beds are an outlier 
group so an exact age is uncertain (Dulhunty & 
Eadie, 1969). They consist of 'chert containing 
Jurassic plant and fish fossils ... in soil derived by 
weathering from Jurassic Purlawaugh sediments 
(Dulhunty & Eadie, 1969). The small bivalve 
faima shows a mixture of depositional environ- 
ments, preserving both isolated, singular valves 
and whole animals in aggregations. Modern 
sphaeriids and corbiculinids exhibit such mass 
aggregations after flooding (pers. obs.). 

Fossil hyriids identified herein occurring in the 
Jurassic have an unknown position within the 
Waloon Coal Measures due to inaccuracies in 
locality data. They are badly preserved and are 
isolated throughout the measures. Both valves 
are usually found intact. 

Tliree major units containing unionoids are 
derived from the Eromanga Basin; Winton, 
Griman Creek and Coreena Formations. The 
Griman Creek and Coreena Fonnation, derived 
from the Rolling Downs Group is considered 
Earlv Cretaceous (early-middle Albian) in age by 
Exon Si Senior (1976) and Burger (1986, 1995) 
and consists of a brackish water unit with out- 
crops within the Surat Basin (Exon & Senior, 
1976). This unit is comprised generally of lithic 
glauconitic sandstones, siltstones and mudstones. 
The yoimger Winton Fonnation, in central and 
southwestern Eromanga Basin, is of latest Albian 
to Cenomanian age (Senior el al., 1978), 
consisting for the most part of lacustrine and 
fluviatile siliciclastic sediments that produced 
labile sandstones, siltstones and mudstones. 

All three units contain mass aggregations of 
bivalves with whole animals preserved in situ. 
Mass aggregations of hyriids occurring today 
tend to be observed after Hooding, hence the 
Griman Creek, Winton and Coreena Fomiations 
may be considered to be more fiuviatile than 
brackish-marine. The presence of marine 
molluscs in two of the formations (Griman Creek 
& Coreena) may then be attributed to marine 
incursions onto the fluviatile systems. As there is 
very poor, if any, recorded stratigraphy the 
occurrence of the taxa in sympatry cannot be 
analysised. The relative abundance, however, 
may give clues to the placement of former 
drainage systems and in what direction they 



408 



MEMOIRS OF THE QUEENSLAND MUSEUM 



flowed. For example, the Winton Formation has 
no record of marine taxa, whereas the Griman 
Creek and Coreena formations have. The Griman 
Creek formation specimens seem to be 
dominated by hyriids and the Coreena formation 
dominated by marine taxa. Examination of all the 
available species and their analogous environ- 
ments need to be assessed in greater detail before 
such conclusions can be evaluated. 

PALAEOECOLOGY 

The families of freshwater bivalves present in 
the Mesozoic (Hyriidae, Glauconomidae and 
?Sphaeriidae) illustrate analogous faunal as- 
semblages to those occurring in modem Australia 
ireshvvater and cstuarine systems. Regardless of 
climate dynamics, one large and several small 
bivalve groups have continually dominated the 
freshwater and estuarine bivalve faunas since the 
Triassic. The large bivalve fauna is comprised 
entirely of the hyriids, and the small bivalve 
faunas of either; glauconomids, sphaeriids or 
corbiculids. During the Mesozoic, there is an 
apparent faunal change occurring at the end of the 
Triassic, early Jurassic, and within the Jurassic 
toward the Cretaceous. 

At the end of the Triassic there were two major 
faunal groups, the large Prohyria and Megalo- 
virgus hyriids and the small Unionella and 
Protovirgus glauconomids. Both these faunas 
have been preserved separately in the fossil 
record and reveal no sign of overlap into either 
faunal system. It is, therefore, probable that the 
small glauconomids inhabited environments not 
suited for the larger unionoids and visa versa. 
Indeed, today glauconomids are only known 
from estuarine environments, and are therefore, 
quite distinct from the freshwater hyriids. This 
provides good evidence that the Wianamatta 
Shales were derived from such a system of 
deposition. 

An analogous system is found today in the 
brackish to estuarine dwelling sphaeriids and 
corbiculids with the modem intercontinental 
riverine hyriid radiation. One can then speculate 
at the life habits of the Mesozoic taxa. The heavy 
shelled, large hyriids were presumably living in 
more lacustrine, Ouviatile regions of Australia's 
Mesozoic, whereas the smaller valved 
glauconomids and ?sphaeriids would have been 
respectively confined to the brackish, estuarine 
and riverine regions of coastal Mesozoic 
Australia. 



SYSTEMATIC PALAEONTOLOGY 

HYRIIDAE Ortmann, 1911 
VELESUNTONAE Iredale, 1934 

Prohyria McMichael 1956 

TYPE SPECIES. Pw}wiajohnstom{E^Qv\di%Qh, 1892); 
liom the Oligocene, West Taniar River. 

DIAGNOSIS. (After McMichael, 1956) 'Medium 
sized to large freshwater mussels of the 
subfamily Velesunionae, the anterior end mod- 
erately to markedly swollen, the posterior end 
drawn out into a bluntly rounded rostration, 
which is of maximum length at a position in the 
middle of the height of the shell; hinge well 
developed, with large cardinal teeth.' Rostration 
of the posterior end is distinct in all growth forms. 
Escutcheon broad, extending almost the entire 
length of the valves. Anterior adductor muscle 
scars deep, elongate, orientated antero-ventrally 
and relatively small. Umbones consistently 
eroded. Prominent hinge with one elongate 
lateral tooth and large cardinals. 

REMARKS. The genus is placed within Vel- 
esunionae due to the absence of beak sculpturing. 
McMichael (1956) erected this genus for the 
placement of two species, P johnstoni, from the 
Oligocene of Launceston Basin and P eryensis, 
from the Late Triassic of Leigh Creek, SA and 
Ipswich Coal Measures, SE Qld, within the 
velesunionines. Ludbrook (1961) assigned 
another large unioid from the Late Triassic, Unio 
springfleldemis, considered as Prohyria spring- 
fieldensis here, from the Springfield Basin, SA. 
Recently, an additional form, P. macmichaeli, 
was described by HocknuU (1997) from the 
Cretaceous of Qld, however, new specimens 
attributable to this taxon from Lightning Ridge, 
NSW, suggests alliance with the hyridellines. All 
three taxa are distinct from any other velesun- 
ionine genus by their large size, thick ornamented 
shell, rostrate posterior profile and distinctly 
inflated umbones. 

Prohyria eyrensis (Etheridge Jr, 1892) 
(Fig. 2K) 

Unio eyrensis Elhridge Jr 1892: 389, pi. 28(1); 1941: 11, pi. 
3(1-3). 

Prohyria eyrensis (Etheridge Jr) McMichael 1957: 228. pi. 

13(8, ll) (non9. 10). 
Unio springfieldensis Ludbrook, 1 96 1 : 145, pi. 2( 1 ,2). 

MATERIAL. AMF51624, 51651, 51642, 51647, 51644, 
51648, 51649, 51625, 51626, 51638, 51640. SAM15473, 
15474. 



MESOZOIC BIVALVES FROM AUSTRALIA 



409 



AGE AND DISTRIBUTION. Late Triassic Irom the 
Black Stone, Tingalpa, Bundamba formations, SE Qld. 
Leigh Creek Formation and Springfield Coal Basin, SA. 

DIAGNOSIS. Large, elongate-oval, equivalved 
unioid with distinct rostration in the posterior 
end. Anterior end inflated and rounded. Hinge 
distinct with strong escutcheon produced along 
most of the dorsal margin. Lunule large and 
excavated. Beaks high and unsculptured. usually 
eroded. Commarginal ornamentation with coarse 
growth lines. Shell thick. From Ludbrook(1961); 
*hinge with two triangular pseudo-cardinal and 
one long posterior lateral in the right valve, one 
triangular pseudo-cardinal and two long 
posterior laterals in left valve' . Anterior adductor 
muscles deep, elongate-ovai and orientated 
anlero-ventrally. 

DESCRIPTION. Shell equivalved, inequilateral, 
umbones inflated and anteriorly placed 1/3 from 
anterior end. Beaks unsculptured but usually 
eroded. Shell thick with coarse growth lines 
producing ridged ornamentation. From 
Etheridge Jr (1892); 'Hinge line straight, ventral 
margin rounded , passing rather sharply upwards 
into the anterior and posterior margins. Anterior 
end small, somewhat acutely curved, posterior 
end narrow, obtusely pointed'. Anterior adductor 
muscle scars deeply set, behind beaks, 
elongate-ovoid and postero- ventral ly orientated. 
Large cardinal teeth, with long bifurcating lateral 
tooth. Morphometries given in Table 1. 

REMARKS. P. eyremis is one of the most 
common taxa found in the Ipswich and Leigh 
Creek deposits. It is a cosmopolitan taxon and is 
the largest in the Mesozoic. Attaining sizes over 
1 2cm long it is comparable to modem species of 
Velesunio and Alathyria. These modem genera 
have been restricted to coastal river systems and 
lakes and may provide good analogues to the 
depositional environment of the Triassic coal 
deposits of SE Qld and SA. 

P. eyrensis was reviewed by McMichael ( 1 956) 
and placed within Prohyria when he erected the 
genus to fit one other taxon, P. johmtoni. Its 
placement within Prohyria is due to the large 
size, wedge-shaped profile, inflated beaks 



positioned anteriorly and the prominent hinge. 

Prohyria eyrensis differs from Prohyria 
johnstoni in its size, size proportions and amount 
of tapering. Prohyria johnstoni tapers more 
sharply to the posterior than Prohyria eyrensis. 
Prohyria springfieldensis is markedly similar to 
Prohyria eyrensis and is synonymised here; 
however, it differs having: finer comarginal om- 
amentation; more rostrate appearance in occlusal 
view; slraighter, more prominent hinge line. P. 
eyrensis differs from Hyridella macmichaeli 
formally placed within Prohyria by Hocknull 
( 1 997) by being much larger, with a more rostrate 
posterior profile. The beak sulcus is relatively 
less produced in P. eyrensis. Juvenile forms of 
eyrensis do not possess the characteristic 
V-shaped beak sculpture, as seen in Hyridella 
macmichaeli. 

On inspection of the holotype and comparison 
to a series of specimens referrable to P eyrensis, 
including the holotype, from the Leigh Creek 
Basin and Ipswich Coal Measures, there is little 
evidence for the separation of these specimens 
fi-om what is obviously a cosmopolitan taxon, P 
eyrensis. 

Prohyria skepticsi sp. nov. 
(Fig- 3) 

TYPE SPECIES. Prohyria johnstoni (E^eridge, 1892). 

ETYMOLOGY. For the Australian Skeptics. 

MATERIAL. HOLOT\TE: UQF52158. PARAT\TES: 
UQF52160, UQF29708A^. 

AGE AND DISTRIBUTION. Jurassic, Waloon Coal 
Measures, Warwick District, SE Qld. 

DIAGNOSIS. Medium-sized, equivalved 
velesunioninc hyriid. Elongate-ovoid, thick shell 
with rugose comarginal ornamentation. Shell 
tapering to the posterior to produce a triangulate 
wedge-shaped posterior margin. Umbones 
placed anteriorly and orientated antero-Iaterally 
to the shells' anterior-posterior axis. Anterior 
adductor muscle scar small, rounded and placed 
high, underneath the beaks. Beaks not sculptured. 
Lateral teeth simple, comprising a single tooth 
and socket in at least the left valve. Hinge line 
long and robust. 



TABLE I . Morphometries (in mm) for Prohyria eyrensis (Etheridge Jr). 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


85.75 


46.57 


34.09 


15.85 


41.04 


16.65 


46.78 


s.d. 


18.11 


10.41 


7.53 


4.02 


9.28 


3.69 


7.75 


Number 


12 


12 


12 


12 


12 


12 


12 



410 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 2. A-C, Megalovirgiis clellandi Holotype QMF29473; A, right valve xO.5; B, dorsal view xO.44; C, 
anterior view x 0.5. D-F, Megalovirgiis wintonensis Holotype QMF34635; D, right valve X 0.7; E, dorsal view 
X 0.7; F, anterior view X 0,8. G-J, Megalovirgiis jaenschi Holotype SAMF 1547a; G, dorsal view X 0.46; H, 
right valve X 0.46; I, anterior view x 0.44; J, right valve x 0.56. K, Prohyria eyrensis Holotype SAMF15473, 
right valve xO.7. L, Alathvria coatsi Holotype SAMF 15477, right valve xO.8. M, Alathyria jaqueti, 
QMF 103903 right valve X T.O. 



MESOZOIC BIVALVES FROM AUSTRALIA 



411 




FIG. l.Prohvria skepticsi sp. nov.: A, UQF52 158 right 
valve xl.2; B, UQF52LS8 dorsal view X 1.0; C, 
UQF29708 left valve xl.5. 



DESCRIPTION. Medium-sized, elongate-ovoid 
hyriid with rugose comarginal ornamentation. 
Shell thick with long thick ligament. Hinge broad 
near umbones and tapers gradually to posterior 
margin. Umbones placed anterior at about 1/5 of 
total length. Posterior margin rounded. Anterior 
margin rounded and begins slightly below beak. 
Beaks not sculptured but possess fme lines 
conspicuous around margin. Escutcheon incon- 
spicuous. Anterior adductor muscles placed 
antero-ventrally to anterior edge of beak. Muscle 
scar small and ovoid. Dentition simple 
comprising single lateral tooth in the left valve 
and possibly two cardinal teeth. Rugose 
ornamentation interspersed by fine growth lines. 
Umbones orientated anteriorly to lateral axis of 
valves. The shell immediately posterior to 
umbones is inflated and produces a slight ridge 
that runs the length of the shell to the posterior 
margin. The ventral is straight and rounded to 
both ends. 



REMARKS. The absence of sculpture and the 
elongate-ovoid shape places this hyriid within the 
velesunonines. It's placement within Prohyria is 
warranted due to the rugose and thick shell inter- 
spersed with fine growth Unes, triangulate-ovoid 
outline of the posterior margin, small ovoid 
highset anterior adductor scar and anteriorly 
placed umbones with laterally orientated beaks. 
Prohyria skepticsi differs from Prohyria eyrensis 
and Prohyria Johnstoni, by it's smaller size, more 
elongate outline, relatively smaller adductor 
muscle, and relatively broader hinge line. 
Morphometries given in Table 2. 

Alathyria Iredale, 1934 

Alathyria jaqueti (Newton, 1916) 
(Fig. 2M) 

Unio jaqiieti Newton, 1916: 230, pi. 6(2-6). 
Velesmio jaqueti (Newton) McMichael, 1957. 

MATERIAL. QMF10S87, 15812, 103859, 103862, 
103868, 103879, 103880, 103892, 103894, 103898, 
103899, 103903. 

AGE AND DISTRIBUTION. Uppermost Albian - 
Cenomanian Formations, Winton, (3riman Creek and 
Coreena. Localities: QML229, QML379, QML570, 
Lightning Ridge and White Cliffs. 

DIAGNOSIS. Large velesunionine hyriid, 
elongate-ovoid with distinct winged dorsal 
posterior rim. Ventral margin of valves in lateral 
aspect show distinct concave form. Beaks 
posterior, relatively inflated. Anterior margin 
tapers sharply from mnbonal region. Umbonal 
region always eroded in large individuals. Long 
indistinct lateral tooth with small indistinct 
cardinals of unknown number 

DESCRIPTION. Shell large, elongate, medium 
sized, equivalved. Shell width narrow in young 
individuals, inflating as the shell becomes larger 
Anterior margin tapers away sharply from beak, 
to a rounded profile. Dorsal edge subparallel with 
ventral margin with a truncated posterior 
producing a 'winged' profile. Beaks relatively 
anterior placed 1/3 from the anterior end. 
Pronounced inflation just postero-ventral of 
umbones candying down to the postero-ventral 
edge. Umbones characteristically eroded with 
some being broken off post-deposition. Beaks 
unsculptured. Anterior adductor muscle scars 
inconspicuous, being small and just anterior to 
the cardinal leeth. Hinge relatively strong and 
short. Ventral margin concave toward the mid 
region. The concavity is more pronoimced in 
larger individuals. Posterior adductor muscle 



412 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. Morphometries (in mm) for Prohyria skepticsi sp. nov. 



UQF No. 


Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Hinge Length 


UQF52I58 


52.41 


30.14 


23.84 


13.35 


28.67 


10.35 


32.32 


UOF5:i60 


43.65 


25.64 


■1 


6.04 


18.23 


•1 




UQF29708A 


48.59 


29.75 


15.90 


11.13 


26.72 


10.09 


27.64 



scars indistinct. Cardinal teeth typically unionid. 
Lateral teeth obscured. Escutcheon narrow and 
encompasses most of the hinge area. Ligament is 
prominent within this area. Morphometries given 
in Table 3. Shell allometry is illustrated in Figure 
8. Note that the specimens allied to this taxon are 
distinct from Megalovirgus wintonensis on the 
basis of height lo length ratios. 

REMARKS. Newton (1915) placed this taxon 
within Unio, as with most other unionoids at that 
time. McMichael's reititerpretation of the 
material lead him to place the taxon within 
Velesunionae, as Velesunio jaqneti. The present 
review agrees with McMichael's placement 
within the velesunionines, based on the absence 
of any distinct umbonal sculpturing in small 
individuals. The shells are also less quadrate than 
those seen most prevalently in the hyridellines. 
The large size, winged posterior and tapering 
anterior margin show^ close similarities to 
Alathyria. The current study has examined many 
more specimens and the overall size shows that 
the taxon shares more similarities to Alathyria 
than Velesunio. A. jaqueti differs from it's 
sympatric taxa by possessing a unique height to 
length ratio (only similar to H. macmichaeli). It 
differs from H, macmichaeli because it is winged 
posteriorly and does not possess juvenile beak 
sculpture. 

Alathyria coatsi (Ludbrook, 1961) 
(Fig. 2L) 

Protovirgiis coatsi Ludbrook 1961 : 146, pi. 2(6). 
MATERIAL. SAM15477. 

AGE AND DISTRIBUTION. Early Cretaceous (Neo- 
comian), Blythesdale Group. Localities: as in Ludbrook 
(1961). 

DIAGNOSIS. Medium-sized, equivalved, in- 
equilateral hyriid with fine comarginal 
ornamentation. Umbones relatively deflated. 
Dorsal margin convex, with ventral margin 
highly concave m lateral profile. 

DESCRIPTION. From Ludbrook (1961); 'Shell 
of medium size, compressed, elongate ... dorsal 
and ventral margins slopes approximately 



parallel; dorsal margin gently convex and 
elevated posterior to beaks then curving more 
sharply downwards to the posterior margin ... 
Posterior ridge fairly well marked and gently 
arcuate. Beaks flattened, apparently un- 
sculptured, not prominent; ligament moderately 
prominent, no lunule visible'. 

REMARKS. Ludbrook placed this taxon within 
Protovirgus with no direct characteristic defining 
it's position within the genus. Inspection of the 
holotype allowed the present study to place P. 
coatsi within the extant genus Alaythria. This 
placeinent is on the basis of the following 
characters present in P. coatsi and Alaytharia: 
convex dorsal and concave ventral margins; 
flattened beaks in most members of the group; 
and marked postero-dorsal wing. The specimen 
differs from other species o{ Alaytharia by it's 
smaller size, more pronounced convexo-concavo 
lateral profile and less prominent winged 
postero-dorsal face. 

HYRIDELLINAE Iredale, 1934 

Megalovirgus gen. nov. 

TYPE SPECIES. Megalovirgus jaenschi (Ludbrook), 
from the Upper Triassic, Leigh Creek Formation, SA. 

ET\'MOLOGY Megalos. Greek, [:)ertaining to the taxons 
lai^e size; -vhgiis for the genus Vvgus which it resembles 
closely. 

DIAGNOSIS. Shells medium to large, elongate, 
knife-like, with umbones inllated and positioned 
extremely anterior. Anterior adductor muscle scar 
set on platform, oblong, ventro-laterally orient- 
ated. Shell thin with little or no ornamentation. 
V-shaped scuplturing in the umbones with 
sculpture proceeding down a postero-dorsal ridge 
in smaller individuals. Ornamentation lost in most 
adult valves. Posterior musculature unknown. 
Escutcheon long and broad with distinct ligament. 

DESCRIPTION. Shells that are distinctly 
hyridelline in juvenile morphology, losing this in 
the larger individuals. Medium to large shells, 
equivalved, inequilateral, elongate with beaks 
placed extremely anterior. Posterior margin 
tapering moderately to sharply, all knife-like. 



MESOZOIC BIVALVES FROM AUSTRALIA 



413 



TABLE 3. Morphometries (in mm) for Alathyria jagueti (UGv/ton). 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


61.82 


34.2 


23.57 


17.09 


34.28 


11.12 


27.44 


s.d. 


12.49 


7.73 


5.67 


4.96 


7.73 


3.40 


6.72 


Number 


12 


12 


12 


9 


12 


7 


3 



Anterior margin truncated and rounded, with 
indistinct lunule, producing weak beak sulcus. 
Fine comarginal growth hnes, with smooth orn- 
amentation. Anterior adductor muscle elongate 
and placed high in the shell, in front of beaks; 
orientated postero-ventrally or just ventrally. 
Raised on small platform. Dorsal and ventral 
margins run subparallel producing elongate 
tear-shaped outline to valves. Escutcheon broad, 
housing distinct hinge with prominent ligament. 
Teeth comprise of one large lateral tooth, weakly 
preserved and two weak cardinal teeth. Typical 
unionid teeth. Valves produced laterally to 
provide distuict height. 

REMARKS. This genus is erected to house 
species that have been confused with another 
family (the glauconomids) of freshwater bivalves 
inhabiting other areas at similar times. The 
Glauconomidae, presented later, has an unusual 
presence within Australia and has evolved to fill 
an ecology not used by unionoids. The species 
formally placed within Protovirgus by 
McMichael (1956) are now placed within 
Megalovirgus except for the type species, P. 
dimstani. This is because all of the other taxa are 
believed to be hyriids, whereas, P. dunstani, is 
thought to be a glauconomid. The reasons for this 
will be presented under P dimstani. 

Megalovirgus differs from all other hyriid 
genera on the basis of the following differences: 
elongate profile with indistinct beak sculpture 
(cf Cucumerunionae); indistinct lunule and weak 
sulcus; fine growth lines with no ornamentation; 
and distinct anterior adductor muscle, raised on 
platfomi, just in fi-ont of beaks. The new genus is 
now used for M. jaenschi, M. femingi, M. 
cleUandi and M wintomrisis. Protovirgus coatsi 
Ludbrook, 1961, is redescribed above as, Alathyria 
coatsi. 

Megalovirgus jaenschi (Ludbrook, 1961) 
(Fig. 2G-J) 

Protovirgus jaenschi Lvidbxook, 1961: 145 pi. 2(3-4). 

MATERIAL. HOLOTYPE: S.4MI547aand PARATYPE 
1 547b. AMF5 1 627, 5 1 639, 5 1 645, 5 1 650, 5 1 63 5, 5 1664. 

AGE AND DISTRIBUTION, Late Triassic, formations: 
Leigh Creek, Springfield Coal Basin. Localities: Leigh 



Creek Coal Fields, Leigh Creek Golf Course, Springfield 
Coal Fields, SA. 

DIAGNOSIS. Large elongate, inflated hyridel- 
line with fine comarginal omamentation. Shell 
thin and valves deep. Posterior edge tapering to a 
relatively pointed end. Anterior rim rounded and 
short. Beaks placed at I/IO the length from the 
anterior end. 

DESCRIPTION. See Ludbrook (1961), Morph- 
ometries given in Table 4. 

REMARKS. M jaenschi is distinct from the 
other species within this genus due to the place- 
ment of the beaks, narrow fonn, overall size and 
orientation of the anterior adductor inuscle scars. 
M jaenschi is longer than M. wintonensis with 
beaks placed more anteriorly. The anterior 
adductor muscle scar is set more ventrally in M 
jaenschi than in M wintonensis. The valves are 
relatively narrower than in M wintonensis. When 
compared to M cleUandi, M. jaenschi is shorter, 
wider with less height. The anterior adductor 
muscles are placed higher in M clellandi. M, 
femingi is smaller, naiTower with thicker oma- 
mentation. Beaks are placed more posteriorly 
than in M jaenschi. 

Megalovirgus clellandi (Hocknull, 1994) 
(Fig. 2A-C) 

Protovirgus clellandi \\oc]sX\\x\l 1994: 146 (fig. lA-D). 

MATERL\L. HOLOTYPE: QN4F29473; 29475. 

AGE AND DISTRIBUTION. Late Trassic, fomiations: 
Black Stone, Tingalpa, SE Qld. Localities: Ebbw Vale and 
Tingalpa, SE Qld. 

DIAGNOSIS. Large, equivalved, elongate hyriid 
with infiated umbonal area and fine comarginal 
omamentation. Hinge straight and long with es- 
cutcheon broad and distinct. Tapering posterior 
to a very pointed posterior margin. Anterior 
adductor muscle scars raised on platfomi just 
anterior to beaks and lunule. 

DESCRIPTION. See Hocknull (1994). 

REMARKS. M clellandi differs from M.jaenchi 
in h's larger size, higher placed adductor scars, 
more tapering posterior and more anterior 



414 



MEMOIRS OF THE QUEENSLAND MUSEUM 




■FIG. 4. A-F, HyridcUa (Protohyridetla) goondiwindit 'fisis\ A-C , .AMF 1 03S4*-J, A. leli vjivc X 1 3. B, ciorstil view 
X 1. 4. C, right valve x 1 .3: D-F. I lok)lvpcQMF5684, DJeft valve 1 .4. F, anterior viev\ x 2.0, F. dorsal view 
X 1.4. G-1., ilyridella (Hyriddlai macmkhaeli Paialype QMF34637, G, right valve x 1.3, H, dorsal view 
xO.9, 1, anterior view x 1.5. HolotypeQMB34636, J, dorsal view xl,S,K, leftvaJve x 1.5, L, anterior view 



MESOZOIC BIVALVES FROM AUSTRALIA 



415 



TABLE 4. Morphometries (in mm) for Megalovirgiis jaenschi (Ludbrook). 





Length 


Height 


Width 


Beak Height 


Beak Length 


Beak Width 


Ligament Length 


Mean 


85.93 


30.51 


34.89 


27.21 


9.53 


13.9 


44.49 


s.d. 


9.51 


7.91 


1.75 


5.78 


2.61 


2.04 


8.03 


Number 


10 


10 


10 


10 


10 


10 


10 



umbonal-beak region. P. jaenschi also has a 
greater height to length ratio. M clellandi is 
longer, narrower, tapers more sharply, has a more 
anteriorly placed beak and has a larger height to 
length ratio than h4. wintonemis. M. flemingi is 
much smaller and the umbones are placed more 
posteriorly than in M clellandi. 

Megalovirgus wintonensis (Hocktiull, 1997) 
(Fig. 2D-F) 

Protovirgus wintonensis Hocknull, 1997: 223 (fig. lA-E). 

MATERIAL. HOLOTYPE: QMF34635. PARATYPES: 
QMF568 1-5682, 34645, 34646, 34634, 34644, 34647, 
34648.AMF15815, 47175, 68358, 103838, 103841, 103843, 
103844, 103847, 104848, 103850, 103852, 103857, 103867, 
103873, 103893, 103897, 103901, 103911. 

AGE AND DISTRIBUTION. Latest Albian - Ceno- 
manian, fomiations: Winton, Grimai Creek, Coreena 
Formation. Localities: OML229, L379, L570, Lightning 
Ridge, White Cliffs, NSW. 

DIAGNOSIS. Medium-sized hyriid, equivalved, 
inequilateral, slightly inflated umbones and fine 
comarginal ornamentation. Hinge straight; 
anterior adductor muscle raised on platfonn. 
Tapering strongly to the posterior with short well 
rounded anterior end. Umbones 1/15 length of 
shell from anterior end. 

DESCRIPTION. Medium to large hyriid with 
produced posterior end. Elongate with umbones 
inflated. Umbones 1/15 length from the anterior 
end. Anterior margin rounded. Hinge straight, 
ligament long and thick. Dorsal margin long and 
straight tapering to a knife-like posterior profile. 
Ventral margin gracile, tapering sharply to meet 
posterior edge of dorsal margin. Fine growth 
lines producing fine comarginal ornamentation. 
Hinge teeth long and thin, cardinals short and 
indistinct, distinctly unionid. Beaks usually 
eroded or small, closely set. Escutcheon long and 



narrow. Lunule short and broad. Anterior muscle 
scars elongate and orientated antero-ventrally. 
Posterior muscle scar unknown, presumably 
small and indistinct. Juveniles possess V-shaped 
rugae on their beaks, being eroded later in life. 
Rugae also occur along the margins of the 
escutcheon. Shell relatively thin. Juveniles start 
out being triangular in profile and becoming 
more elongate produced shell. Morphometries 
given in Table 5. AUometry given in Figure 8. 

REMARKS. M wintonensis differs from M 
clellandi in having a smaller profile, wider and 
thicker shell, less anterior umbones, narrower 
escutcheon and less tapering at the posterior end. 
M jaenschi is larger, has a less developed 
anterior adductor muscle scar which is placed 
lower in the valve than in M wintonensis. M. 
fenungi\s smaller, with the umbones less anterior 
than in M wintonensis. 

Hyridella (Hyridella) Swainson, 1840 

Hyridella (Hyridella) macmichaeli 

(Hocknull, 1997) 
(Fig. 4G-L) 

Prohvria macmichaeii Hocknull. 1997: 224 (fig. 2A-C), (non 
fig. IF-H). 

MATERIAL. HOLOTWE: QMF34636. PARATYPES: 
5677, 34637, 34638. Additional Material. AMF68346, 
103864, 103866, 103869, 103870, 103871, 103874, 
103878, 103882, 103884, 103886, 103889, 103890, 
103910. 

AGE AND DISTRIBUTION. Cenomanian, fomiations: 
Winton Fomiation (type) and Griman Creek. Localities: 
QML379, OML570, QML229 and Lightning Ridge. 

DIAGNOSIS. 'Medium-sized, equivalved 
unioid, ovate, rugose ornamentation, umbones 
anterior and inflated, beak slightly sculptured, 
shell thick' Hocknull (1997). Juveniles with 
V-shaped ornamentation. Anterior muscle scar 



TABLE 5. Morphometries (in mm) for Megalovirgus wintonensis (Hocknull). 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


51.83 


22.7 


16.42 


11.63 


20.76 


7.10 


20.34 


s.d. 


20.98 


9.71 


9.19 


5.7 


9.10 


3.72 


8.5 


Number 


20 


20 


20 


18 


19 


17 


14 



416 



MEMOIRS OF THE QUEENSLAND MUSEUM 



set below beak and raised slightly on platform. 
Simple unionid teeth, with two peg-like cardinals 
and one long lateral tooth. 

DESCRIPTION. Medium-sized, elongate-ovoid 
unionid. Equivalved with slightly inflated um- 
bones. Hinge line distinct and convex, tapering to 
the posterior to produce a pointed posterior 
profile. Beaks 1/5 from the anterior end, usually 
eroded. Fine growth lines produce ridged 
commarginal ornamentation. Thick shell. 
Escutcheon broad and ligament short and 
distinct. Juveniles have V-shaped rugae on the 
beaks, absent in the adults as the beak is eroded. 
Anterior adductor muscle scars small and set 
below the beaks, raised on platforms. Unionid 
teeth with a long lateral tooth and small indistinct 
cardinals. Morphometries given in Table 6. 
Allometry shown in Figure 8. 

REMARKS. The presence of V-shaped rugae as 
beak sculpturing in smaller individuals places 
this taxon within the hyridellines and more 
specifically Hyridella. The anteriorly placed, 
inflated and thick shelled umbones coupled with 
tapering elongate-ovoid profile ally P. 
macmichaeli to Prohyria. The taxon is smaller 
than the other members of this small but cosmo- 
politan genus. The type species, P. johnstoni 
(Etheridge Jr, 1892) of McMichael (1956) is 
longer, wider and the dorsal margin produces a 
sharper, tapering edge. P. eyrensis is also longer, 
wider with the umbones placed fiirther anterior. 
Unio springfieldensis has similar attributes as P. 
eryemis, being longer, wider with anteriorly 
placed umbones. This taxon also tapers more, as 
in P. johstoni. It seems that Hyridella shares many 
similarities with Prohyria; however, this is 
apparently due to convergence and not homology. 

Hyridella (Protohyridella) Cotton & Gabriel, 

1932 

Hyridella (Protohyridella) goondiwindiensis 

(Hocknull, 1997) 
(Fig. 4A-F) 

Velesimio goondiwindiemis Hockiiull. 1997: 225 {fig. 2D-J) 

MATERIAL. HOLOTYPE: QMF5684, 5683. PARA 
TYPES: 5685, 5686, 34639-36341 AMF68342, 103849, 



103853, 103858, 103865, 103881, 103883, 103895, 
103896, 103900, 103902, 103904, 103906, 103907. 

AGE AND DISTRIBUTION. Latest Albian - Ceno- 
maniati, formations: Griman Creek Formation (type) aiid 
Coreena Fonnation. Localities: Goondiwindi (type), 
Liglitning Ridge and Wliite ClilFs. 

DLAGNOSIS. Small, unioid with fine comarginal 
omamentation. Quadrate-angulate with strong 
postenor ridge. Umbones relatively inflated and 
positioned anteriorly. Beaks sculpturing, 
V-shaped in juveniles, eroded in adult forms. 
Teeth simple and unionoid. Anterior adductor 
muscle small placed just in front of beaks. Shell 
expanded posteriorly, forming a slight winged 
appearance in adult fonns. 

DESCRIPTION. Small, equivalved quadrate- 
angulate hyriid. Fine growth lines fonn ridges 
producing distinct commarginal omamentation. 
Beaks distinct and usually eroded laterally. 
Umbones anterior, situated 1/3 from the anterior 
margin. Ligament short and distinct producing a 
strong hinge. Hinge line tapers smoothly to a 
rounded posterior margin. Antero-ventral margin 
distinctly convex. Shell relatively thick. 
Juveniles with distinct V-shaped rugae on the 
beak, lost in the adult. .luveniles are more ovoid 
without the pronounced posterior ventral margin 
of adults. Lunule small and indistinct. Es- 
cutcheon broad with the borders smooth and 
tapering to the posterior. Anterior adductor 
muscle scars small, oval and just beneath the 
beak. Teeth simple unionid. Morphometries given 
in Table 7. Valve growth pattern is illustrated in 
Figure 8. 

REMARKS. Initially, Hocknull { 1 997) placed //. 
goodiwindiensis within Velesiinio due to it's 
overall similarities in morphology. However, 
Hyridella {Protohyridella) goodiwindiensis is 
distinctly hyridelline because of the presence of 
distinct V-shaped beak sculpture in the juvenile 
forms (McMichael & Hiscock, 1958), now 
known from additional specimens. Placement in 
Hyridella is somewhat tentative, due to the 
marked difference in maximum sizes and relative 
sizes, when compared to modem taxa of the same 
genus. However, the overall variation within 



TABLE 6. Morphometries (in mm) ior Hyridella (Hyridella) macmichaeli (Hocknull). 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


43.15 


26.04 


19.52 


11.79 


23.64 


8.2! 


14.80 


s.d. 


15.74 


9.80 


7.28 


4.94 


8.75 


3,34 


6.74 


Number 


14 


14 


13 


14 


14 


12 


11 



MESOZOIC BIVALVES FROM AUSTRALIA 



417 



TABLE 7. Morphometries (in mm) for Hyridella (Protohyridella) goondhvindiensis (Hocknull). 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


28.6 


18.68 


14.36 


8.55 


18.61 


6.55 


10.39 


s.d. 


4.48 


3.87 


2.23 


L79 


3.82 


1.46 


3.31 


Number 


14 


14 


14 


14 


14 


14 


14 



Hyridella supports a conservative approach to 
higher taxonomy in this group (McMichae! & 
Hiscock, 1958) and therefore placement within 
Hyridella is warranted. When compared to 
modem Hyridella species it is evident that this 
taxon is much smaller with finer growth and 
thinner shell and is, therefore, distinct from them. 
It does, however, bear striking similarities to the 
type species of the monotypic subgenus Proto- 
hyridella. The small size, quadrate-angulate fonn 
with posterior ridge prominent is characteristic of 
Hyridella (Protohyridella) glenelgensis (Dennant). 
Extinct Hyridella species are also a lot larger than 
this taxon, with the exception ofPalaeohyridella 
godthelpi gen. et sp. nov., suggesting that this 
species is a dwarf representative of the hyridel- 
lines leading towards Protohyridella. Wlielher 
this provides evidence for separation of 
Protohyridella as a distinct genus is controversial 
when dealing with unioid species concepts 
(McMichael& Hiscock, 1958). When comparing 
the allometry seen in H (P) goondiwindiensis 
with that of Palaeohyridella godthelpi gen. et sp. 
nov. it can be shown that, even though they 
develop similarily sized adults, the growth 
pattern to this end is quite different, being 
positive allometry in H. (P.) goondiwimiiensis 
and negative allometry in Palaeohyridella 
godthelpi gen. et sp. nov. 

Hyridella whitecliffsensis (Newton, 1916) 

Unio whUecliffsensis Newton, 1916: 231 pi. 6 (7-8). 
Hyridella whitecliffsensis (Newton), McMichael, 1957: 240. 

AGE AND DISTRIBUTION. Early Cretaceous, fomi- 
ations: Coreena. Localities: While Cliffs NSW. 

DIAGNOSIS. Shell small, equivalve, ovoid 
hyriid of hyridelline affinity. Moderately infiated 
valves with V-shaped beak sculpturing. 'Periodic 
growth divisions, and mnnerous, close-set, 
microscopical concentric striations ... posterior 
ridge become angulate' (Newton, 1916). 

DESCRIPTION. See Newton (1916). 

REMARKS. The holotype of this taxon was un- 
available for study during this review, however, it 
is possible to diagnose this taxon from other 
members of Hyridella. The most apparent 



distinguishing features being the angulate 
posterior margin and concentric growth striae. It 
differs from H. (Protohyridella) goondiwindi- 
ensis by it's smaller size and more angulated 
posterior profile. It differs from Palaeohyridella 
godthelpi gen. et sp. nov. by it's smaller size, less 
pronounced posterior ridge, closer growth 
ornamentation, more ovoid profile and narrower 
shell. 

Hyridella (Protohyridella) sp. 

(Eig. 5A-C, D-E) 



MATERIAL. UQF52157, UQF52159, 
UQF44278, U0F44276, UQF44277. 



UQF5216 



AGE AND DISTRIBUTION. Jurassic, Waloon Coal 
Measures, Warwick District and Rathdowney area, SE 
Qld. 

DESCRIPTION. Medium-sized, equivalved, 
ovoid hyriid with the comarginal ornamentation. 
Beaks eroded. Posterior margin tapers abruptly at 
the posterior 2/5 of valve, producing a Iri- 
angulate-ovoid posterior margin. Small anterior 
adductor muscle scar placed antero- laterally to 
lunbone. The escutcheon is shallow. The anterior 
margin begins 2/3 of the beak height and 
produces a small rounded margin about 8. 19mm 
to the anterior. The dorsal margin is inflated distal 
to the anterior and is inflated past the imibones, 
tapering to the posterior. This produces a wedge- 
shaped hinge jutting out of the general line of the 
shell. The umbones are placed anteriorly to the 
general line of the valves. The ventral margin is 
rounded to both ends. Ligament short. L)QF52157 
& 52161 are both medium-sized inequivalved, 
ovoid inflated hyriids. The inflation and 
inequilateral morphology is interpreted as an 
abnormal growth fonn for the shells. The general 
morphology follows that described above. 
Morphometries given in Table 8. 

REMARKS. The material is placed within 
Hyridella (Protohyridella) based on the follow- 
ing characteristics: 1 , beak sculptured and eroded; 
2, inflation posterior to beak and producing a 
flanged ventral margin; 3, shell thin with fine 
comarginal ornamentation. Due to the paucity of 
specimens and closeness in morphology to 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIQ 5. A-C, Hyridella {Pm(oltyri<i$I{ci) sp, UQFS^l^SJ A, right valye xO.8; B, dorsaf view X i:0; C anterior 
view X0,7, D-E, deformed ifldivJ<tial of ffyrtd^ta (Protohyridella) sp. UQF52I61;D, left valve KOi; E, 
dorsal view X 0.8. F-^yPeUaaofrjfrid9l^^dthelpig^. pt^jy.jiov.; F-l, M, HoloQipe AMF 1039 12; F, dar^^l view 
X 2.0; G, right valve x2.(j; IT, VffLve k 1 .4; I. dors^-raesj^ ><, K4; J, left valve X 1*4; L, anterior viB^v X Lfi. 
K-li,>I^Paraty{ie-AMPlD39i3; K, letl valve x J.4; L,^disal! view ^i^iN, aritftSailview x U6, 



specific assignraent is not warranted until more 
^chitens -are ^vajlaUe. 

Mes^Ofayridella McMiph^el, 1^56 

rVPE SPECIES. M ipsviciems (EUieridge Jr,l892) from 
the Upper TViassic, i^^cli Coal MeBsures. 



DIAGNOSIS: Sx&ati ihgirWdlitte Wioid with 

elotigate-o\'al. equivalvcd, inequilateral valves. 
Beaks anteriorly placed with small anterior 
adductor muscle scar, ovoid and set anteriorly to 

produced, widely concave. Mod9t^)^iii€ate4 



MESOZOIC BIVALVES FROM AUSTRALIA 



419 



TABLE 8. Morphometries (in mm) for Hyridella (Protohyridella) sp. 



UQF No. 


Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Hinge Length 


UQF52157 


37.76 


28.95 


? 


9 


27.54 




7 


UQF52159 


45.46 


31.24 


23.65 


8.17 


28.79 


10.83 


27.36 


UQF52161 


50.73 


35.26 


7 


? 


3 1 .90 


15.03 


28.34 



REMARKS. McMichael (1956) erected this 
genus to place one species of small ?hyridelline, 
M ipsviciensis. It's affinities with modern 
subfamilies is thought to be hyridelline, however, 
this is superficial. Unfortunately, no more mat- 
erial has shed any light on its relations within the 
hyriids. 

Mesohyridella ipsviciensis (Etheridge Jr, 
1892) 

Vnio ipsviciensis Etheridge Jr, 1892: 388 pL 42(2-3). 
Mesohyridella ipsviciensis (Etheridge Jr) McMichael, 1957: 
238 pL 8(5). 

AGE AND DISTRIBUTION. Late Triassic, fonnations: 
Black Stone, Tingalpa, Bundainba, SE Qld; Springfield 
Coa! Basin. Localities: Ebbw Vale, BundaiTiba, Tingalpa, 
Bundamba, SE Qld, Springfield Coal Fields. 

DIAGNOSIS. Shell small, elongate-oval with 
little winged posterior. Shell thick with strong 
ornamentation. 

DESCRIPTION. From McMichael (1956); ^Small 
freshwater mussels of uncertain affinity, but pos- 
sibly belonging to the subfamily Hyridellinae. 
Shells elongate-oval, not winged, moderately 
swollen. Dorsal margin behind the beaks more or 
less straight, then curving rather sharply 
downwards, and descending obliquely to fonn a 
rather blunt posterior end with the ventral 
margin. Beaks not elevated or swollen, heavy 
corroded, sculptural characters unknown. Shell 
surface marked with fairly strong growth lines. 
Hinge characters and muscle scars unknown'. 

REMARKS. Etheridge Jr (1892) described M 
ipsvicensis within Unio. McMichael (1956) 
proposed a genus for the species, in the absence 
of the holotype, and called it Mesohyridella. The 
present study has not located the holotype either, 
however, on inspection of many more unionoids 
recovered from the Ipswich Coal Measures (the 
type locality is within this fonnation) other spec- 
imens are allocated for this species. The hyriids 
that resemble the descriptions of Etheridge Jr and 
McMichael have strong ornamentation, relatively 
uninflated umbones and beaks placed more 
central than P. eyrensis, the other large ovoid 
hyriid present in the fauna. This morphologically 



distinct group of bivalves are present in the 
Blackstone fonnation, Tingalpa formation of the 
Ipswich Coal Measures, SE Qld and the Spring- 
field Basin, SA. 

Palaeohyridella gen. nov. 

TYPE SPECIES. Palaeohridella godthelpi gen. et sp. 
nov. from uppennost Albian - Cenomanian Giiman Creek 
Formation, Lightning Ridge. 

ETYMOLOGY. Palaeos, Greek, representing the ancient 
aspects of the taxon and -Iryridella tor the affinity to the 
freshwater bivalve genus Hyridella. 

DIAGNOSIS. Small hyriid unioid of the 
hyridelline subfamily. Umbones inflated and 
placed extremely anterior, 1/13 of total length 
from anterior end. Posterior margin quadrate- 
angulate with prominent umbones to posterior 
ridge. Ridge preserving prominent ridging, 
developing into V-shaped sculpturing toward the 
beaks. Escutcheon broad, relative to length. 
Ornamentation coarse with fine comarginal 
growth lines. Anterior profile truncated and 
rounded, producing weak beak characteristics. 

DESCRIPTION. Small, equivalved, inequi- 
lateral, hyriid with inflated umbones. Umbones 
greatly anterior being less than 1/13 the length 
from the anterior. Anterior profile truncated and 
rounded. Posterior end produced, producing a 
tear drop valve shape. Shell thick, growth lines 
fine, producing ridged comarginal ornament- 
ation. Ventral margin gracile and convex. Hinge 
line straight with short thick ligament. Escut- 
cheon broad and distinct with ornamentation 
producing a postero-dorsal ridge extending down 
both valves to the posterior end. V-shaped rugae 
on beaks and along postero-dorsal ridge on 
juveniles and adults. The rugae being present on 
the ridges in adults only. Beaks distinct and 
close-set. Lunule very short but relatively broad. 
Muscle scars unknown. 

REMARKS. This genus is erected due to the 
presence of a taxon unlike any hyriid described 
previously. The V-shaped beak sculpturing in the 
juvenile and adult allies the new genus to the 
hyridelline lineage (McMichael & Hiscock 
1958). When compared to Hyridella (Hyridella) 



420 



MEMOIRS OF THE QUEENSLAND MUSEUM 



spp. and Hyridella (Pwtohyridella) spp. there are 
marked dilTerences; these are I, urnbones more 
anteriorly situated; 2, broad, distinct escutcheon; 
3, posteriorly rostrate profile; and 4, truncated, 
convex anterior margin. 

Palaeohyridella godthelpi gen. et sp. nov. 
(Fig. 5F-N) 

ETYMOLOGY. For Henk Godthelp, a good friend who 
brought some of the specimens to my attention. 

MATERIAL. HOLOTYPE: AMF103912; 26 Paratypes. 

AGE AND DISTRIBUTION. Uppemiost Albian - Ceno- 
manian, formations: Griman Creek Fomiation. Locality: 
Lightning Ridge. 

DIAGNOSIS. As for genus. 

DESCRIPTION. As for genus. Morphometries 
given in Table 9. Allometry given in Figure 8. 

GLAUCONOMIDAE Gray, 1853 

Glauconomids are a group of estuarine to 
brackish water bivalves from the east coast of 
Australia, inhabiting mangrove systems, 
mudflats and sands. There are four extant species 
recognised from Lamprell & Healy (1998); 
Glauconome plankta, G. virem, G rugosa and G 
cerea. 

Unionella Etheridge, 1879 

TYPE SPECIES. Unionella wiancmuittemh Etheridge Jr, 
1888 from Late Triassic, Gibraltar Tunnel. 

DIAGNOSIS. Small, elongate-oval glauconomid. 
Beaks anterior and close-set. Umbonal region 
slightly inflated devoid of sculpturing. Posterior 
margin slightly winged in larger individuals. A 
series of characteristically elongate beak muscle 
scars is present just below the beak commissure. 
Hinge short and indistinct. Anterior adductor 
muscle scars deep, elongate and usually placed 
anteriorly to umbones. Muscle scar triangular in 
lateral profile. Two to three cardinal teeth 
present, median and posterior most distinct. 

REMARKS. The small size, ovoid shape, deep 
adductor muscle scar and winged posterior along 
with the 'clumping' nature of the taxa suggest that 
this genus has evolutionary affinities to the 



modem Glauconomidae, and certain morph- 
ological features similar in Anthracosiidae. 
Anthracosiids have a well known presence in the 
Carboniferous and Permian of Eurasia. 
McMichael (1956) in his review of these forms 
suggested such an affinity of Unionella with the 
anthracosiids. When comparing these shells to 
the modern Glauconome spp. there are marked 
similarities: valves are closely set with the beak 
not well produced; a series of small beak muscle 
scars occur under the lip of each beak; shell is 
produced anteriorly, directly in ft-ont of the beaks, 
compared to on the midline as seen in the 
anthracosiids; tAvo to three small cardinal teeth, 
increasing in size from the anterior end. The 
median and posterior cardinals are largest and are 
close-set. 

This genus was described for three apparently 
distinct forms, Unionella howralensis, U. wiana- 
mattensis and U. carnei. Comparing the overall 
morphological diversity in species from an- 
thracosiids and glauconomids it is apparent that 
Etheridge Jr ( 1 882) did not take into account the 
possible phenotypic plasticity. I have clumped 
the three forms into one species, as I believe there 
is not enough consistent variation to propose 
different species. The taxon available for this is 
U. wianamattensis. 

Unionella wianamattensis Etheridge Jr 1 888 
(Fig. 6K) 

Unionella hawralemis Etheridge Jr, 1888: i3, pi. I & 2, figs 

8-14; McMichael, 1956: 236, pi. 13, fig 6. 
Unionella carnei Etheridge 1888: 14, pi. 2, figs 5-7; 

McMichael, 1956: 237, pi. 13, fig. 7. 

MATERIAL. LECTOTYPE: AMF35775; 20183, 21085, 
35769, 35771. 3987, 20184. 35766, 35764, 35778, 35773A, 
3577B, 

AGE AND DISTRIBUTION. Late Triassic, Formations: 
'transition beds between the 1 lawesbuo" Sandstone and the 
Wianamatta Group' (McMichael, 1956). Localities: 
Gibraltar Tunnel, Bowral, Smith's Brick Quarry at Crown 
Street, Waterloo and Surrey Hills, NSW. 

DIAGNOSIS. Shell small, equivalved, elongate- 
ovate, with fine comarginal ornamentation. 
Valves inflated mid-laterally, however, umbones 
relatively flattened. Shell slightly winged 
posteriorly. Anterior adductor muscle scar deep. 



TABLE 9. Morphometries (in mm) for Palaeohyridella godthelpi gen. et sp. nov. 





Length 


Height 


Width 


Beak Length 


Beak Height 


Beak Width 


Ligament Length 


Mean 


21.88 


15.5 


12.47 


3.65 


14.39 


4.12 


9.21 


s.d. 


2.59 


1.99 


2.56 


0.62 


2.5 


0.79 


1.42 


Number 


15 


15 


15 


15 


15 


15 


15 



MESOZOIC BIVALVES FROM AUSTRALIA 



421 




FJG 6. A-F, h'otovir^us hrook\'uIcrtsi.\ sp. nOV.; A-B, Paralvpe AMF41440, righl valve x3.0. C-D. Paralype 
AMF4I442. \ci\ valve xMt. H-F. ilololype AMFI9773", left valve X3.0. G-J, Pmfovirgiis dtinsfanil Q 
AMF356'>3, righl valve >: 2.0. M. AMF35776, leA valve showing beak sculpliire (bs) and cardinal leeth (cl) 
X 1.6» I, AMF35693, right valve showing beak sculpture X 2.25, J, AMF35776,dorsal view showing,cardinal 
teeth L7. K. Umnelia wiammattensis AI^STTS. rjiB^ Vilve x3.0. 



422 



MEMOIRS OF THE QUEENSLAND MUSEUM 



elongate and postero- ventral ly orientated. A 
series of beak muscle scars extend just postero- 
dorsally of the anterior adductor scar. 

DESCRIPTION. From Etheridge (1888): ^Shell 
ovate-obliquely oblong, laterally compressed, 
thin. Dorsal margin or hinge line straight 
posteriorly, angulated at the anterior end, but in 
its entire length not as long as the shell; ligament 
small, and projecting but little above the dorsal 
margin. Ventral margin nearly straight, with a 
slight sinus at the middle. Anterior end small, 
very much compressed, and with the margin 
rounded; posterior end compressed, the diagonal 
ridge well marked. Although not strong; the 
flanks of the valves decrease rapidly in convexity 
from this ridge to the ventral margin, but an 
almost imperceptible sinus traverses them 
upwards from the ventral marginal inflections'. 
Forms initially described as Unionella 
bowralensis and Unionella carnei the shell 
shows more inflation in the valves. Wlien height 
to length was measured for the specimens 
described by Etheridge Jr (1888) a consistent 
allometry is illustrated. 

Morphometries are presented for Unionella 
bowralensis^ Unionella camel and Unionella 
wianamattensis in Table 10. 

REMARKS. The three species initially decribed 
within Unionella are here synonymised as one 
species on the basis of the following similar but 
variable characteristics: elongate-ovoid shells 
that have similar growth allometries; slight 
winged postero-dorsal margin; and sympatric 
occurrence in the same horizon. As alluded to in 
McMichael (1956)'s review of Unionella, the 
overall shell variation in anthracosiid taxonomy 
is well documented and provides evidence that 
these three taxa are one. Glauconomids are also 
highly variable. 

Protovirgus McMichael, 1956 

TYPE SPECIES. Protovirgiis dumtani (Etheridge Jr) trom 
the Upper Triassic Wianainmatta Shale, Sydney Basin. 

DIAGNOSIS. Small, elongate, equivalved, 
inequilateral glauconomid with long tapering 
posterior profile. Umbones flattened and placed 
extremely anterior. Deep, long, posteroventrally 
orientated anterior adductor muscle scars. A 
series of beak muscle scars placed just below the 
beak commissure. Three cardinal teeth, median 
and posterior cardinal largest and closely spaced. 
Reduced or no lateral tooth. 



TABLE 10. Morphometries (in mm) for Unionella 
wianamattensis Unionella bowralensis) 
(Etheridge Jr), Unionella wianamattensis 
(^Unionella carnei) (Etheridge Jr), Unionella 
wianamattensis (Etheridge Jr). 



AMFNo. 


Length 


Height 


Width 


Beak 
Length 


Unionella wianamattensis (^Unionella bowralensis) 


20183 


21.7 


8.75 


5.15 


6.8 


20185 


12.4 


7.7 


5.6 


3.05 


35769 


12.15 


6.7 


5.4 


3.15 


35771 


9.2 


5.7 


4.15 


2.15 


Mean 


13.86 


7.212 


5.075 


3.78 


Unionella wianamattensis (^Unionella carnei) 


3987 


12.45 


6.7 


4.05 


3.75 


20184 


12.55 


7.7 


5.9 


3.1 


35764 


18.35 


9.6 


7.5 


5.9 


35766 


18 


8.6 


7.7 


5.1 


35778 


16.7 


8.55 


6.05 


4.7 


Mean 


15.01 


8.23 


6.24 


4.51 


Unionella wianamattensis 


35773a 


15.4 


8.7 


5.65 


4.9 


35773b 


12.15 


8.75 


5.4 


3.55 


35775 


14.8 


8.8 


4.5 


3.85 


Mean 


14.11 


8.75 


5.183 


4.1 



REMARKS. This genus is related to Unionella in 
the possession of beak muscle pits, deep, 
elongate anterior adductor muscle pits and 
slightly wider postero-dorsal margin and three 
small cardinal teeth. The genus was described by 
McMichael (1956) to accommodate two species, 
R dimstani and P, flemingi, from the Late Triassic 
of Sydney and Cretaceous of New Zealand, 
respectively. Unfortunately, P. flemingi was not 
examined during this study and will not be 
considered here. 

Protovirgus dunstani (Etheridge Jr, 1888) 
(Fig. 6G-J) 

Unionella dunstani V.X\\QX\AgQ h 1888: II pi. Ull-IQ). 
Protovirgus dunstani (Etheridge Jr) McPvlichael, 1957: 232 
pi. 14(8). 

MAFERIAL. LECTOTYPE: AMF35693; 35776, 35777, 
35870. 

AGE AND DISTRIBUTION. Late Triassic, fomiations; 
'ti-ansition beds between the Hawesbuiy Sandstone and the 
Wianamatta Group' (McMichael, 1956). Localities: 
Gibraltar Tunnel, Bowral, Smith's Brick Quarry at Crown 
St, Waterloo, and Suney Hills, NSW. 



MESOZOIC BIVALVES FROM AUSTRALIA 



423 




F[G 7. A-B, Prolosphaeiium gianae sp. nov.; A, AMF38165 Hololype, right valve x4.0, B, AMFl03934b 
Paratype, left valve x 5.0. C-D, Protosphaerium talbragaremis gen. el sp. nov.; C, AMF103934a Hoiotype, 
right valve x4.0, D, AMF103929 Paratype right valve x5.4. 



DIAGNOSIS. Small, equivalved, glauconomid. 
Elongate shell, tapering to posterior end. Beaks 
anterior, with flattened umbonal region. Beaks 
insignificant and closely set. Postero-dorsal 
margin slightly winged. Three cardinal teeth; 
large median and posterior cardinal and small 
anterior cardinal, no lateral tooth. 

DESCRIPTION. From Etheridge (1879), ^Shell 
narrow, very transversely elongate, thin and 
compressed throughout its length ... cardinal 
margin very long and slightly arched . . . anterior 
end very much compressed, the margin rounded, 
posterior end thin, attenuate ... umbones placed 
close to the anterior end, small, and laterally 
flattened ... anterior adductor impression fan- 
shaped, situated very high up under the anterior 
cardinal margin; umbonal scars veiy strongly 
marked, two immediately behind adductor scars 
in a line 

Morphometries given in Table 1 1 . 

REMARKS. P. dimstani is unique within it's 
fauna, differing from other members of 
Unionellahy the extremely elongate nature of the 
valves. The presence of beak scars illustrates 
genetic relation to UnioneUa, however, it differs 



so markedly from them that separation at the 
generic level is relevant. P. diwslcmi difl'ers irom 
P. flemingi by it's smaller size and less tapering 
profile. The only other species of this genus is 
Protovirgiis brooh^alensis sp. nov. which differs 
in being smaller and more rounded posteriorly. 
The anterior adductor muscle in P. brookvalensis 
sp. nov. is much longer and situated higher than P. 
diifistan'L The umbones are more flattened in P. 
dimstani than in P. hroo/mi/ensis sp. nov. 

Protovirgus brookvalensis sp. nov. 
(Fig. 6A-F) 

ET^'MOLOGY. For tlie type locality, Brookvale quany. 

AGE AND DISTRIBUTION. Late Triassic, formations: 
Wiananialta Shale. Localities: Brookvale Quany, NSW. 

IVIATERL\L. HOLOTYPE: AMFI9773. PARAT^TES: 
19805, 43401, 41438, 41442, 41440, 41439, 49805. 

DIAGNOSIS. Small, equivalved glauconomid 
with umbones anteriorly placed. Elongate valve 
shape with tapering posterior edge. Winged dorsal 
margin produced by ridge emanating from the 
umbonal area and terminating toward the 



424 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE IL Morphometries (in mm) for Protovirgus 
dunstani (Etheridge Jr). 



AMFNo. 


Length 


Height 


Width 


Beak Length 


35693 


34.7 


11.9 


? 


8.85 


35776 


37.2 


12.2 


6.05 


8.65 


35777 


25.8 


11.8 


4.8 


5 


35870 


40 


16.6 


8.75 


9.05 


Mean 


34.42 


13.12 


6.53 


7.88 



posterior end. Umbonal region anterior with 
beaks closely set. 

DESCRIPTION. Holotype F19773, is a small 
equivalved, inequilateral, elongate-ovoid 
anthracosiid. Umbones anteriorly placed at 1/5 
from the anterior end. Beaks weak with no sculp- 
turing. Hinge line straight and long producing a 
straight dorsal profile. Rounded posteriorly. 
Anterior end rounded. Muscle scar unknown. 
Teeth unknown. Subumbonal ridge runs postero- 
ventrally away trom the timbo. Fine comarginal 
growth lines with weak ornamentation. 
Morphometries presented in Table 12. 

REMARKS. The elongate nature of the valves, 
anteriorly placed umbones and tapered posterior 
end place this taxon firmly within Protovirgus. 
Morphological features that differentiate P. 
brookvalensis from P. dunstani, the only other 
member of this genus, are it's smaller size, 
weakly defined umbones, more rounded 
posterior end, weaker anterior muscle scars. 

7SPHAERIIDAE Jefferys, 1862 

Protosphaerium gen. nov. 

TYPE SPECIES. Protosphaerium talbragarensis gen. et 
sp. no\'. Jurassic, Talbragar Fossil Fish Beds. 

ETYMOLOGY. Proto, pertaining to this being the first of 
it's kind, -sphaerium, for the genus Sphaerium, 

DIAGNOSIS. Small, equivalved, ovoid bivalve 
with beaks subccntral. Fine comarginal growth 
lines with rugose comarginal ornamentation. Hinge 
short and convex in lateral profile. Ventral margin 
rounded to fonn distinct 'pea shelF like shape. 

DESCRIPTION. Small, equivalved, equilateral 
with fine growth hnes. Little or no ornament- 
ation. Shell very thin with umbones just slightly 
inflated. Ovoid with beak 1/3 length from anterior 
end. Posterior profile rounded continuous with 
convex hinge outline. Anterior profile also 
rounded with a small insignificant lunule. 
Escutcheon narrow and also insignificant. Hinge 
short with hinge ligament relatively thick. 



TABLE 12. Morphometries (in mm) for Protovirgus 
brookvalensis sp. nov. 



AMFNo. 


Length 


Height 


Width 


Beak Length 


43401 


22.5 


9.85 




9.85 


41438 


23.15 


9.95 


9 


10.7 


19773 


25 


10.5 




12.1 


41442 


23.8 


8.75 


7 


11.6 


41440 


22.4 


8.7 


n 


4.45 


41439 


21.5 


9.2 


7 


10.55 


19805 


20.7 


9.1 


9 


8.6 


Mean 


22.72 


9.43 




9.69 



Muscle scars unknown, presumed weak. Teeth 
unknown or insignificant. 

REMARKS. The pea-shell shaped valves, sub- 
central beaks, small size and clumping deposition 
post death, are all characteristics of members of 
the Sphaeriidae (Kuiper, 1 983 ) and Corbiculidae. 
As this genus is smaller than those normally 
found within the corbiculids it is concievable that 
this taxon belong within the sphaeriid lineage. As 
the tooth morphology of the type species is un- 
known it is with some degree of uncertainty that 
this placement within the Sphaeriidae is given. 

Protosphaerium talbragarensis gen. et sp. nov. 
(Fig. 7C-D) 

ETYMOLOGY. For the type locality of Talbragar. 

MATERIAL. HOLOTYPE: AMF103934A. PARA- 
TYPES: AMFl 03929, 103935(b), 59823. 

AGE AND DISTRIBUTION. .lurassic, formations: 
Talbragar Fossil Fish Beds. Localities: Talbragar. 

DIAGNOSIS. As for genus. 

DESCRIPTION. As for genus. Morphometries 
given in Table 13. 

REMARKS. This taxon bears marked affmhies 
to the small fi'eshwater bivalves from the family 
Sphaeriidae. Uncertainty surrounds the decision 
to place Protosphaerium talbragarensis within 
the Sphaeriidae, however, it seems plausible that 
this group has a Jurassic presence in Australia 
and has remained here since then. Since it fits in 
no other family and the erection of a new family 
seems unwarranted, it is here placed within the 
sphaeriids. 

Protosphaerium gianae sp. nov. 
(Fig. 7A-B) 

ETYMOLOGY For Gian Holmes. 



MESOZOIC BIVALVES FROM AUSTRALIA 



425 



TABLE 13. Morphometries (in mm) for Proto- 
sphaerium talbragaremis gen. et sp. nov. 



AMFNo. 


Length 


Height 


Width 


Beak Length 


103929 


6.9 


4.3 


7 


2.9 


103934 


11.5 


6.8 


? 


4.25 


103935b 


10.75 


6.25 


? 


4.2 


59823 


10.45 


6.15 


? 


4.15 


Mean 


9.9 


5.87 





4.93 



TABLE 14. Morphometries (in mm) fox Protosphaerium 
gianae sp. nov. 



AMF No. 


Length 


Height 


Width 


Beak 
Length 


38165 


9 


6.25 


? 


3.7 


103931 


5.7 


4.1 


? 


2.5 


103934B 


6.55 


4 


9 


3.1 


103932 


7.65 


4.5 


9 


3.1 


Mean 


7.22 


4.71 





3.1 



MATERIAL. HOLOTYPE: AMF38165. PARATYPES: 
AMF103931, 103932, 103934B. 

AGE AND DISTRIBUTION. Jurassic, formations: 
Talbragar Fossil Fish Beds. Localities: Talbragar. 

DIAGNOSIS. Small, equivalved, inequilateral 
freshwater bivalve of possible corbiculid affinity. 
Fine conmarginal growth lines and weak 
ornamentation placed periodically along the valves. 
Characteristically winged toward the posterior, 
producing a quadrate lateral profile to the valves. 
Beaks closely set and anteriorly placed. Hinge 
long with one thin, indistinct lateral tooth. 
Cardinals and anterior laterals unknown. 

DESCRIPTION. Small, equivalved, inequilateral 
with fine growth lines. Subtriangular. Some 
periodic ornamentation produced by thickenings 
in the shell. Winged posterior produced from 
ridge originating from the umbonal region. 
Posterior end produced and somewhat rostrate. 
Anterior profile roimded. Beaks weak, closely set 
and placed anteriorly. Hinge weak with one thin 



O) 

c 



X 

> 

0) 



0.9 
0.8 
0.7 
0.6 

0.5 
0.4 
0.3 
0.2 H 
0.1 




indistinct lateral tooth. Cardinals and anterior 
laterals unknown. Muscle scars unknown, pre- 
sumed weak. Morphometries given in Table 14. 

REMARKS. Placed tentatively within Proto- 
sphaerium, this species shares some characters 
with Protosphaerium talhragarensis. Until more 
material presents itself, this genus provides an 
adequate position. In some respects, Proto- 
sphaerium talhragarensis sp. nov. is similar to 
Batissa (Batissa) violacea (Lamark) within the 
corbiculids. While similar in morphology this is 
not considered as having close genetic affinity. 
Proto. gianae is much smaller and with reduced 
hinge length. 

ACKNOWLEDGEMENTS 

I thank the Malaeological Society of Australia 
for their financial assistance in the form of a 
grant. The Australian Skeptics are thanked for 
their ongoing support. Henk Godthelp and the 
University of New South Wales is thanked for the 
provision of new material. Alex Cook, Natalie 








20 



80 



100 



40 60 
Total Length (mm) 

FIG. 8. Bivarate plot of relative height/length, plotted against length for hyriids from the Cretaceous. • = 



Palaeohyridella godthelpi gen. et sp. nov., ^ Hyridella (Hyridella) macmichaeli, * 
hyridella) goondiwindiensis, o = Protovirgus wintonensis, a = Alaytharia jaqueti. 



Hyridella (Proto- 



426 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Camilleri and Paul Tiemey are thanked for their 
assistance and provision of workspace. Kevin 
Lamprell, Daryl Potter and John Stanisic are 
thanked for their assistance and provision of 
comparative modern specimens. Robert Jones 
and Neville Pledge are thanked for the provision 
of fossil specimens from the Australian and 
South Australian Museums respectively. Mary 
Wade and Ralph F. Molnar are thanked for the 
collection of most of the Cretaceous specimens. 

CITED LITERATURE 

BURGER, D. 1986 Palynolog\', cyclic sedimentation 
and palaeoenvironments in the Late Mesozoic of 
the Eromanga Basin. Geological Society of 
Australia: Special Publication 12: 53-70. 

BURGER, D. 1995. Timescales. 9. Cretaceous. AGSO 
Record 1995/38:1-35. 

CHAPMAN, F. & COOKSON, l.C. 1 926 A revision of 
the 'Sweet' Collection of Triassic plant remains 
from Leigh's Creek, South Australia. Trans- 
actions of the Ro\ al Society^ of South Australia. 
50: 163-178. 

DE JERSEY, N.J. 1975. Microspore zones in the Lower 
Mesozoic of southeast Queensland. Pp. 159-163. 
In Campbell, K.S.W. (eds) Papers from the Third 
Gonwana Symposium. (ANU Press: Canberra). 

DE JERSEY, N. J. & HAMILTON, M. 1965. Triassic 
mircofloras of the Mooroka and Tingalpa 
Formations. Queensland Government Mining 
Journal 66:315-318. 

DULHUNTY & EADIE, 1969. Geolog> of the Talbragar 
Fossil Fish Bed Area. Journal and Proceedings, 
Royal SocieK of New South Wales, 102: 1-4. ^ 

ETHERIDGE Jr, R. 1888. Small unio fauna from the 
Wianamatta Shales. Memoirs of the Geological 
Survey NewSouth Wales, Palaeontology 1 12-16. 
1892. The organic remains of the Ipswich Form- 
ation. In Jack, R.L. & Ethridge, R. (eds) The 
geology and palaeontology of Queensland and 
New Guinea. Geological Survey of Queensland 
Publication 92. 

EXON, N. & SENIOR B.J., 1 976 The Cretaceous geology 
of the Eromanga and Surat Basins. BMR Journal 
1,33-50. 

HAAS, F 1969. SuperramilyUnionacea.PpN41 1-468. 
In Moore R.C. (ed.) Treatise on invertebrate 
palaeontology Part N. Vol. 2, Mollusca 6, 
BivaK ia. (Geological Societ> of America and 
Universit^ of Kansas: Lawrence, Kansas). 

HILL, D., PLAYFORD, G & WOODS, J.T. 1965. 
Triassic fossils of Queensland. (Queensland 
Palaeontographical Society: Brisbane). 

HOCKNULL, S.A. 1994. A new Triassic frehwater 
bivalve from southeast Queensland. Memoirs of 
the Queensland Museum 37( 1 ): 1 46. 



1 997 Cretaceous Freshwater Bivalves from Queen- 
land. Memoirs of the Queensland Museum 
42(1): 223-226. 
JOHNSON, W. 1960. Exploration for coal, Springtield 
Basin, in the hundred of Cudlamudia, Gordon- 
Cradock District. Rep. Invest. Geological Survey 
of South Australia 16:62. 

K.UIPER, J.GJ, 1983. The Sphaeriidae of Australia. 

Basteria47: 3-52. 
LAMPRELL, K.L. & HEALY, J.M. 1998. Bivalves of 

Australia Vol. 2. (Backhuys Publishers: Leiden, 

The Neatherlands). 

LOVERING. J.F. & ELROY, C.T 1969. Wianamatta 
Group. Pp. 417-423. In Packham, GH. (ed.) The 
geology of New South Wales. Vol. 16 Part 1. 
(Geological Society of Australia). 

LUDBROOK, N.H. 1961. Mesozoic non-marine 
Mollusca (Pelecypoda: Unionidae) from the north 
of South Australia. Transactions of the Royal 
Society of South Australia. 84: 139-148. 

McMICHAEL, D.F. 1956 A review of the fossil 
freshwater mussels (Mollusca, Peleycopoda) of 
Australasia. Proceedings of the Linnean Societ>' 
of New South Wales 81 : 232-233. 

McMICHAEL, D.F. & HISCOCK I.D. 1958. A mono- 
graph of the freshwater mussels (Mollusca: 
Peleycopoda) of the Australian region. Australian 
Journal of Marine and Freshwater Research 9: 
372-507. 

NEWTON R.B. 1915. On some molluscan remains 
from the opal deposits ( Upper Cretaceous) ofNew 
South Wales. Proceeding of the Malacologica! 
Societ>- 10:217-235. 

PARODiZ, 'j.J. & BONNETTO, A. A. 1963. Taxonomy 
and zoogeographic relationships of the South 
American Naiades (Pelycopoda: Unionacea and 
Multelacea). 

PLAYFORD, G & DETTMAN, M.E. 1965. Rhaeto- 
Liassic Plant Microfossils from the Leigh Creek 
Coal Measure, South ./\ustralia. Senck. Leth. 46 
(2/3): 127-181. 

ROGERS, M.J. 1965. A revision of the species of 
nonmarine bivalvia from the Upper Carbonif- 
erous of eastern North America. Journal of 
Paleontology 39: 663-686. 

SENIOR, B.R., MOND, A. & HARRISON, RL. 1978. 
Geology of the Eromanga Basin. Bureau of 
Mineral Resources, Australia, Bulletin 167. 

WADE, R.T 1953. Note on a Triassic fish fossil from 
Leigh Creek, South Australia. Transactions of the 
Royal Society of South Australia 76: 80-81 . 

WEIR, J. 1960. AMonographofBritish Carboniferous 
non-marine Lamellibranchia. Palaeontolo- 
graphical Societ\ : 273-320. 



A^'^<!L^Hm(PO^^VAi DL M()SP()N(;iAE: MICRDCK>^I^I&A^iF»lJ^MTHE 

WJiST£ltN INDIAN OCEAN 



Hopper. I.N. A,. Kellv. M. & Kennedy. I.A, ?(iOO 06 1&: A new Claihno (Porifcro: 
Dttnospangiac: MicrocioniUacJ from the WcsLcm Intliaii Oceah.^fiym/Zrv r;/ //jt {JuLvn-slaml 
Mtt^An^im^^^^^^m^iS^ 0079-8835. 

A nt-w species of microcionid marine sponge Ciuthria {MicrocUma) rkhmondi is^iim. is 
<lescnbcdfrom>Zaazabar, Tanzania, and is higiilv unusual in having exlrcmclj' large accolada* 
tW^JtSi tbftt foi^ ^S^^igfnata and ^^kulclul tracts wilhtn Mic ctiuunosoinal and uctuKomal 

skeretoiut iu^Si^effiasy ^ ecbumting aic^titI)os^4e3) ^ live blue colDurs^tioA mi 9 

species is compared 'WtlnthdtltbfiK'.^j^^ of Clathria desoribt^ fitittif Wtsstmi IttdiQD' 

g»san. Southeast PMa and AfaM^A thllf-Red Sea provinced^ ^dmbtirjp^TeMftK!»V^ 
vc toxodt^mgta. n Pofifero, Demospon^iae, PoecHoseleri^, Micr0ciom<i^^ 

apeciiiS, laxonOfW^K Zanzibca: Western IncHurt Ocean. 

Jofm M.A. N(Mj/Kr ii'muii J(tlinH{aijin-qid.}^oy.uui ami John A- Kcfifmh . Ouccnsialicl 
MusemfJ, !'<) Hox 3300, South Brisbane 4101, Ansimha: Michelle Kell\\ Marine lieolo^' 
and Aquaculture. Nutiomil Insitiute oj Water aihi Atmosphttic Research (NIIVA}, Private 
3ps lf^^9S^ jVcwpTo^M Auckim^ itfw Zeaiandi H Deoemher 1999. 



The extensive literature oa BpongBS of the 
Western Indmn O^ean is evfdeno^ ofa? ncb:m4 
diverse fauna {Table 1). liven thoilgh'tlift 

iiucnlur> iilready cuniuins 'several hundred' 
species (Kelly, 19i)7). ir is undoubtedly far from 
conipleie. For example. Van Soest (!'^"^Mb) 
coUeclcd 240 species &om ihe Seychelles ^xid 
AiinGt^ i^iatids^MVU^as only 1 67 species were 
Knpwn pitvyJusly from fhe region. 

Within the Western Indian Ocean (i.e, 
extending along the eoniinenta! shelf from Nalal 
to Somalia and ineludmji the islands from 
Madagascar to the Seychelles; Richmond, 1997), 
Southeast African regions (Ntff-Jtl to tlw Cape) 
9n4 Msfmx Gulf-Red Sea^provnuscs 74 species 
bf i&ficroclonidae have been te|n>rted (Hooper, 
K»Q6a), of which 64 belong to Clathria, 10 still 
unnamed (Tabic 2). Of these, 44 (or 6^>%) have 
fu.ii yet been (oiind t)uts!de tlie region. This 
number of ^apparent endemics" may decrease 
over time us the regitm'*s InbdiVersUy becomes 
bcttd" known, but it still represents an extraordinarily 
high level of cndcmisra compared to other sponge 
genera and other phyla. For example. Van Soesl 
(i994b) found about 24*,' of all sponges col- 
lected from the Seychelles and Aminnitcfeteft^S 
«(^/9i^inic to the W Indian Ocean fauna, and 
Rldwtoml (I'W?) suggestedih^otjly about 1 5% 
of all the regional marine biota may be endemic 
to the \V Indian Ocean region. 

Biogeograpliic altlnitics ot the sponge launa 
are Iboughi to b€f essentially TethyAn (VSui2065^ 



1994a), with ,southcrn incursions of species of 
Ggndwaiwa origin (Hooper & Levi, J994; 
MbOpen 1996a). This fauna is thought to be moFrt: 
similar to the centred Indian Ocean and Indo- 
Malaya regions (Van SoesI Si llajdu, IQ^^?), and 
inthisregSfdfefllows the general pattern seen in 
other Bi^ri^ Mvertehj-atc ph^a t<;.fi^ scli^i- 
actinarjaii ^SOFafe Vbrtm, I^rVtron, 1993V 
Richmd!^.(]y'?7) suggested fliai about 35"": vf 
the Wesft Indian f)ccan marine biota wa.s widely 
distributed, extending into the Red Sea aitd 
perhaps ajj far as t^e lud^-Malay region, 35% 
ranged across the whole Itido-Pacific region, and 
] 5% extended into the warm temperate region&of 
the Atlantic Ocean, lb date only one niicro- 
etonid. Cluihna ( Thalysiusl \ntlpma (Lamarck), 
appear^i lu be truly In4o-Ptt(afic '$Q$t«10^^1^^% 
with :a confimted' distributidn extending ftdm 
Tongu to Ihe Red Sea (i.e. with conspecific'dy 
cttnlmncd through niorphological comparisons 
heiANeen recent vol lee| tans ofliving populali;<ns, 
liqqDcr, 199(7a}. HQjvcvcru even tlda tii^ding has 
yetiirbctonlTnfled through genetic ftnalysi* to 

delcrmine whether slight nior])hoiogieal ditler- 
enccs between regional populations represent 
intraspecitic variabibtv or indicate the existci'.cc 
of a series of possible allopamc sibling species. 

Van Soest (1994b) also noted a high level of 
regional heterogeneity between sponge faunas of 

the Seychelles and Amirante Island groups, wilh 
only 17% of species conimon to both leginnal 

launas. This supports a simitae finding ^rom 



428 MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 1 . Literature on the sponges of the Western Indian Ocean, Southeast Africa and Arabian Gulf-Red Sea 
provinces. 



Province 


Locality 


Sponge literature 


Coastal East Africa 


Zanzibar 


Lendenfeld, 1897;Baer, 1905; Jenkin, 1908; Burton, 1959; Thomas, 1976a, 1979b; Pulit- 
zer-Finali, 1993; Kelly, 1997; Magnino & Gaino, 1998 


Tanzania 


Jenkin, 1908; Thomas, 1976a; Pulitzer-Finali, 1993; Magnino & Gaino, 1998 


Mozambique 


Levi, 1964; Thomas, 1979a, 1979c, 1980a, 1980b; Laghi et al., 1984; Schmidt et al., 1997 


Kenya 


Marsden. 1975; Bruce, 1976; Thomas, 1981a; Vacelet et al., 1991; Pulitzer-Finali. 1993 


Somalia 


McCabe et al., 1982; Finamore et al, 1983; Hooper, 1996a 


South Africa 


Ehlers, 1870; Carter, 1871; Gray, 1873; Vosmaer, ISSO; Kirkpatrick, 1900, 1901, 1902a, 
!902b, 1903, 1904, 1908, 1913; Sollas, 1908; Stephens, 1915; Burton, 1926, 1929, 1931, 
1933a, 1933b, 1936, 1958; Levi, 1963, 1967; Borojevic, 1967; Day, 1981; Schleyer, 
1991; Pettitetal., 1993b; Rudi et al, 1993, 1994a, 1994b, 1995; Barkai et al, 1996; 
Hooper et al., 1996; Samaai, 1997; Beukes et al, 1998; Koren Goldshlager et al, 1998; 
McPliailet al, 1998 


Offshore East Africa 


Madagascar 


Bosraug, l913;Decary, 1946; Levi, 1956; Vacelet & Vasseur, 1965, 1966, 1971, 1977; 
Vacelet, 1967a, 1967b, 1977; Vacelet etal, 1976; Ivanova et al, 1993; Hooper, 1996b 


Aldabra 


Levi, 1961 


Comoros 


Sara etal, 1993b; Pettitetal, 1993a, 1994a, 1994b 


Reunion 


Levi, 1986; Aknin et al, 1996 


Mauritius 


Thomson, 1868; Duncan, 1880; Topsent, 1890; Dendy, 1922; Van Soest, 1993 


Saya de Maiha 


Dendy, 1922; Kolbasov, 1992 


Seychelles 


Wright, 1881; Ridley & Dendy, 1887; Topsent. 1893a; Dendy. 1922; Levi, 1961; 
Thomas, 1973, 1979c, 1981b; Hooper & Krasochin, 1989; Ngoc Ho, 1990; James etal 
1991; Venkateswarlu etal, 1991; Van Soest, 1994b; Van Soest etal, 1994; Trimurtulu 
& Faulkner, 1994; Hooper, 1996a; Pettitet al, 1997 


Amirante 


Carter, 1880; Ridley, 1884; Ridley & Dendy, 1887; Dendy, 1922; Van Soest etal, 1994; 
Braekman etal, 1998 


Northwest Indian 
Ocean 


Red Sea 


Keller. 1889, 1 89 1 ; Topsent. 1892; Row, 1911; Levi, 1958, 1965; Burton, 1959; Delseth 
etal, 1979; Sara etal, 1979; Kashman etal, 1982, 1989; Mergner, 1982; Sokoloffet al. 
1982; Mebs, 1985; Carmely & Kashman, 1986; Vine, 1986; Gebreyesus et al, 1988; Ilan 
& Loya, 1988, 1990: Carmely et al, 1990; Kolbasov, 1990; Isaacs & Kashman, 1992; 
Rinkevich et al, 1993; Rudi & Kashman, 1993; Kelly-Borges & Vacelet, 1995; Guo et 
al, 1996, 1997a, 1997b; Ramadan, 1997; Beer& Ilan, 1998; Worheide, 1998 


Ethiopia 


Isaacs et al., 1991 


Eritrea 


Hooper, 1996a 


Arabian Sea 


Carter. 1869; Topsent, 1893b; Dendy, 1913, 1915, 1916a, 1916b, 1916c, 1922; Kumar, 
1924a, 1924b. 1^24c, 1925; Burton & Rao, 1932; Burton, 1959; Thomas, 1975, 1976b, 
1979b, 1988. 1989; Rahim, 1979; Kamat et al, 1981; Patel et al, 1985; Kondracki & 
Guyot, 1987; James et al, 1989; Parameswaran et al, 1989, 1992a, 1992b, 1994, 1997; 
Kobayashi et al, 1992a, 1992b; Pettibone, 1993; Sara & Bavestrello, 1995; Bavestrello et 
al, 1996; Thomas etal, 1997 


Oman 


Sara & Bavestrello, 1995; Bavestrello et al, 1996 



Sponge surveys of NW and NE Australian reefs 
(Hooper, 1994; Hooper et al, 1999), with the 
implication that taxonomic inventories of regional 
sponge faunas are largely incomplete, with pos- 
sibly many new taxa remaining to be discovered 
within these highly heterogeneous and 'apparent 
endemic' regional populations. The present study 
describes one such species discovered during 
routine surveys of Zanzibar undertaken by MK as 
part of a project to produce an inventory of the 
marine fauna and flora and popular field guide to 
the region (Richmond, 1997). 

Methods for preparation and examination of 
material are described by Hooper (1996a). 
Spicule measurements refer to (minimum-(mean)- 
maximum) dimensions of lengths and widths taken 
from 25 random samples of each spicule category 



and are given in micrometres unless otherwise 
stated. Abbreviations: BMNH, The Natural History 
Museum, London; QM, Queensland Museum, 
Brisbane. MK is grateful to Dr Matthew Richmond 
for facilitating her participation in the East Africa 
marine storveys. 

SYSTEMATICS 

PORIFERA Grant 
DEMOSPONGIAE Sollas 
POECILOSCLERIDA Topsent 
MICROCIONINA Hajdu, Van Soest & Hooper 
MICROCIONIDAE Carter 

Clathria Schmidt, 1862 

Clathria (Microciona) Bowerbank, 1862 
Refer to synonymy in Hooper (1 996a) 



A NEW CUniRiA FROM Tiffi DSDLVN OCtAN 



43^ 



Claibria (Microciona) i-ichmpnijl.^. tov. 

(Higs 1-4) 

KyMOlOGy..f'ftf prMattl^ Riclmiond, Institute 
oC^^bi^ iS?%aoeSi "Ze^izibar, in nscognition ol' his 

^Itna and Sud^ of E Aflmekf C6adhtt tlticbrhCttKl, 

M^TEiaiiL. iiOU)TYPE. yM<:i306785 (fragineni 
BMNH 1995^29.96). E side of P;Inla^ Sandbank Rwsf; 
Zanzibar Towu Un^uja I.. Taiizmuci. 6" 10.0*5, 39*a3'E, 
10m depth, 7.viii.lW. coil. M. Kelly, SCUBA. 

DISTRIBUTIOM. Known only from (he fringing 
rects ctfTZan^ibur Tovmi, Unguja Island, cnciiisl- 
ing deaU coral i>ubstrale on it shallow ("ringii^g. 
reef. 

DLSCRIPTION. "shape. Very ihinlv riu:rnslmfz 
(p.2-0,7ruiU tliick) in small patches tl<)--20cni 
aiametcT^'drcompldely env eloping ^piilTtlbiM*. 
< ..•luui Ri^vLil l>liic vvitli a ^^iojet tittgc? alive, 
brownish-orange m cthanul. 

Oxcuks. Large (up to iwn diameter); rafeed on 

nu'inhnnciuiN lip (approximately 4nmi hiuli), 
.scattered in or entire surface and w ilh proniinenl, 
V«in-'i^--e,, iLiiJi.d ; iibsurlace drainage canals radi- 
ating tuwarUs cagh oscule; oscules aiKl ^Irmn^^gv 
canals collapsed up^n^eSe^ation. 

sinyolh. llcsliy surface. 

Sdtosome, No sp6i5wl category of ectosonval 

spicules present, altbougb ehoanos(Mnal princip- 
al styles arising from ihc iindt:rlylni.! skeleton. 
Standing perpendicular to tlie suhslraic, may pro- 
trude a long way through llie surlace. Bundles ot 
toxodragmata occasionally lie on ih^ surface, 
although nwi of these appear to be confined 
below iIk peripheral skeleton. 

S!fhi\-/o.sfmic. [iclow the surface are plumose 
hundli;^ p£ayx>Uary subtylostylqs^jnpstlyxunning 
pcqicndieuliii'and paratangentiaHo. oVtfccasioh- 
ally protruding ihiough. the surface. These 
subeclosomal skeleial bundles form stellate 
brashes aasocialcd with (or parallel to) (he lart*er 
pro^fuding choatjosomai principal styles, 
Tbxodragraala fomi thick b untiles If^Iftw the 
peripheral skeleton, resembling jnegasdere 
spicule tracts, lying liuijgtintial to til* sUr&ce dild 
scattered between tSe creel choaHtlGainal 
pnncipal s<ylc», 

CJtoMoifiv^. "fclfdrodrottid skeletal siniclnre, 
with thin hvinedesriMuid basal layer of spongin 
iibre. approxunaiciy 30 iliick. l\ighly collag- 
evums, granul^Tj dm^lc brcms .pigmented, willi 



cafcitic detritus embedded beneath; basal 
Spongin with iifjfcsdv dispmed echihatiAg 
aeanthostyles fewliednBd ^ j,staMding per- 
pendicular to substrate, and 'bttlbous sponL::n 
fibre mides uplo I So thick fotmdonly in ihicUr 
sections of the encrustation: eacii bnlbuns fhrc 
node discrete, erect, without any anaslomo:-^*s 
between adjacent nodes, and each vviih .-5 
choanosomal principal styles embedded and 
perpendicular to subslrate, with spictJlCSs 
diverging sliglitly. becoming plimiose towards 
surface and pioirudtnii up to 350 through 
cctORomc; smaller cchinatin^ aeanihostyhs 
uonfioedniostly (o hyn\cdestnioi^ bas^ Spongin 
tm^r TaFetjr' &een on bulbbU^ fiht^ Elides; 
conversely, principal styles only seen on hutbous 
fibre nodes and not on hymedesinioid basal 
fibics. Dense horizontal bands of accolada 
toxodragmata (up lo 70 (hick) occur abtiul 
taiBWfty throui^h ilic clioaiiosomal skeleton 
cross-section, and also in tlic cctosomal region, 
lying on or below the surface; few jsingle toxa» 
observed in the mesohyl mostly comprismg 
wing-shaped Cotms, whereas most toxas I'ornn'ng 
dragmala. Palmate isochelue moderately 
abundant within mesohyl; collagen within 
mesohyl dense, relatively smooth, nioderatcly 
heavily pigmented oranue-brown; {Shoanoqrte 
chambw*;longaic-oval, up lo 40 X f2. 

A/sga«)t7i?>iK*v Cff<itoo«<mi«1 piihcSfial Rtylcs and 

subtylostylcs very long, slender, slightly curved 
near basal end. long tapering fusifoim points 
base either slightly constricted or non-tylotc. 
smooth or very occasionally with aiusoNCOtc 
lenninations. Length t7fr(463:7>622i \)riAh 
6-(l0.2)-l2. 

Subcctusomal auxiliary subtylostylcs xery 
long. ver\' slender, straight, llisiform points, wi^h 
well di^v^lopcd, entirely amooth subtylote bases, 
tcnglib l9S-t35l.1SV428. widlh 2K3^>5. 

Echinatingacanthostyles relatively uncoiTunou. 
short, slender. 5>ln»ighl ot vciyf slightly cui-ved at 
centre, fusifbrm^-pointed, with moderately well- 
devtfJopcd hasaf coustriction: shaft and base with 
vteStigial graruilwr spines confined mainly to basal 
half of spicule. Lengtti 58-(8i*.6>l34, width 
3-(5.4).8. 

Mkrosrlcre,^. Palmate ^soehcllac fnod6ra<dy 
common, wcU-siticified, with thick, \Ncli- 
dcveli)ped aiac comprising over 7(1% oi spicule 
length. Length 14-(15.2)-17. 

Toxas in two forms: Accolada loxas extremely 
abundant, cxceptiunally long rtnd very ^lende^» 
with iHght ««^«tral curvature an4 ^tcoighl 



430 



MEMOIRS OF THE QUEENSLAND MUSEUM 





FIG 1 . Megascleres from Clathria (Microciona) richmondi sp. nov. (holotype QMG306785). A-B, Choanosomal 
principal st>ie and subtylote base. C-E. Larger echinating acanthostyle, base and shaft, illustrating mostly smooth, 
tubercLiiate ornamentation. F-H, Smaller echinating acanthostyle. base and shaft with rudimentar>' small spines. 
I-J, Subectosomal auxiliar)' subtylostyle and smooth base. 



(non-reflexed) amis, invariably forming toxo- 
dragmata. Length 262-(50L3)-975, width L5- 
(1.9)-2.0. Shorter toxas present but uncommon, 
intermediate between wing-shaped and accolada 
in morphology, with slight to moderate central 
curvature, slightly retlexed arms; found in 
toxodragmata together with accolada toxas and 
also occasionally singly within the mesohyl. 
Length 84-(114.8)-154, width 0.8-(1.04)-L5. 

REMARKS. Clathria (Microciona) richmondi 
sp. nov. is unusual in having 1) huge accolada 
toxas in dragmata, forming dense bands both 
within the mesohyl and lying tangential to the 
surface; 2) a second, less common and much 
smaller form of toxas, intermediate between 
wing-shaped and accolada morphology, scattered 
singly within the mesohyl; 3) a skeleton 
composed of hymedesmioid basal spongin fibres 
in thinner sections and microcionid bulbous 
spongin fibre nodes in thicker parts of the 



skeleton, each node with one or few choanosomal 
principal styles perpendicular to the surface; 4) 
relatively uncommon echinating acanthostyles, 
with rudimentary spination, apparently confined 
to the hymedesmioid basal skeleton; and 5) 
distinctive field characteristics including a royal 
blue colour, large oscules with a prominent raised 
Mip' and prominent subsurface drainage canals 
radiating towards each oscule. Although individ- 
ually these distinctive characters are not unique 
amongst known species of Clathria, in com- 
bination they clearly differentiate the Zanzibar 
species from others. 

1 ) Microsclere morphology, including toxas, 
appears to be a relatively consistent and useful 
character to differentiate between similar species 
(Hooper, 1996a). Six species of Clathria have 
been recorded with accolada toxas forming 
toxodragmata (C. (Thalysias) cactiformis 
(Lamarck), C. (Microciona) densa (Burton), 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



431 





C. (T.) lendenfeldi Ridley & Dendy, C (T.) 
oxitoxa Levi, C. (Axociella) thetidis (Halimann) 
and C. (T.) vulpina (Lamarck) (Hooper, 1996a)); 
refer to Table 2 for known distributions. In C. 
richmondi the accolada toxas are exceptionally 
large compared to most species of Clathria 
(262-975 long), and also form very prominent 
bands within the skeleton. By comparison, those 
of C. (T) cactiformis are 8-355 long, C. (M.) 
dema (205-305), C. (T.) lendenfeldi (7-361 ) and 
C. (T.) vulpina (8-200), and are scattered through- 
out the skeleton in loose bundles not forming 
skeletal tracts. In contrast, accolada toxas of C 
(A.) thetidis and C. (T.) oxitoxa are much larger 
than those of C. (M.) richmondi (175-1280 and 
170-3000 long, respectively), and moreover 
those of C. (T.) oxitoxa also form skeletal tracts 



within the skeleton (Levi, 1963). In this regard C. 
oxitoxa is most similar to C. richmondi, although 
all three taxa differ in virtually every other 
respect (see redescription of C. oxitoxa below and 
C. thetidis in Hooper (1996a)). 

2) Many species of Clathria have two toxa 
morphologies, and this feature is probably of 
little diagnostic importance above the species 
level. Of the species mentioned above only C. 
densa and C. vulpina lack both morphologies of 
toxas. 

3) Hymedesmioid - microcionid skeletal 
structure has been used in the past as a primary 
diagnostic character for several nominal micro- 
cionid genera (e.g. Axocielita de Laubenfels, 
Hymantho Burton, Leptoclathria Topsent), 
although this view is no longer widely held (Van 



432 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Soest, 1984; Hooper, 1996a). 
Within Clathria these species 
are now placed in either the 
subgenera Microciona or 
Thalysias , depending on 
whether ectosomal 
specialisation is absent or 
present, respectively. 
Worldwide there are hundreds 
of encrusting microcionid 
species with hymedesmioid 
and/or microcionid spongin 
fibre skeletons and a mineral 
skeleton composed of perpen- 
dicular and/or plumose spicule 
tracts, of which 22 occur in this 
region (Table 2; species anno- 
tated (2)). 

4) The presence or absence 
of echinating spicules, the 
degree to which they are 
smooth or spined, and the 
morphology of spines have 
been used as generic char- 
acters within Microcionidae at 
one time or another (e.g. 
Anaata de Laubenfels, 
Axociella Hallmann, Folitispa 
de Laubenfels, Isociella 
Hallmann, Ophlitaspongia 
Bowerbank, Paratenaciella 
Vacelet & Vasseur, Temciella 
Hallmann). The absence of 
echinating megascleres re- 
mains a valid diagnostic 
character (at the subgeneric 
level) for some taxa (e.g. 
Isociella^ Axociella^ Ophlita- 
spongia)^ by virtue of the 
consistent combination of this 
feature and the possession of 
unusual skeletal structures 
characterising each of the taxa 
(Hooper, 1996a; Howson & 
Chambers, 1999). Within 
Clathria s.s. there are also 
several species that have lost 
echinating spicules (e.g. C. (C.) pancispicida 
(Burton), C. (T.) craspedia Hooper). By 
comparison, the absence (loss), rudimentary 
development and shape of spines on echinating 
spicules vary widely amongst the many hundreds 
of species of Clathria, although these features 
appear to be consistent at the species level (e.g. C. 
(M) aceratoobtiisa (Carter) with virtually 




FIG. 3. Skeletal structure of Clathria (Microciona) richmondi sp. nov. 
(holot>'pe QMG306785). A, Hymedesmioid-microcionid basal skeleton 
with tracts of auxiliary subt>'lost>ies lying in multispicular bands within the 
choanosome. B, Erect echinating acanthostyles (mostly smooth) per- 
pendicular to the hymedesmioid basal skeleton. 



smooth spicules, C (C.) kylista Hooper & Levi 
with rudimentary spination, C. (T.) dubia 
(Kirkpatrick) with large, heavy spines). Within 
this continuum C. richmondi is most similar to 
the C. kylista condition. 

5) As far as can be ascertained from the liter- 
ature and from personal experience with the 
Microcionidae of the Indo-Paciflc no other species 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



433 



has a deep royal blue colour in life, but the 
importance of this character is limited by the lack 
of good knowledge of their living characters, 
most taxa known only from presei^ved specimens. 
Prominent raised oscules and a subsurface water 
canal system (radiating towards the oscules and 
producing a stellate surface pattern) are features 
common to many encrusting sponges (in which 
the aquiferous system is marginalised onto the 
external surface by virtue of the reduced choano- 
somal thickness), whereas in many microcionids 
this drainage system often has a different 
pigmentation from the adjacent ectosome. 

Of the 64 species of Clathria recorded from the 
Western Indian Ocean, Southeast Africa and 
Arabian Gulf-Red Sea provinces 19 have 
aecolada toxas (Table 2; species annotated (1)). 
Most of these species differ substantially from C. 
richmondi in major features such as growth fomi, 
skeletal structure, spicule geometries, spicule sizes, 
possession of specialised ectosomal skeleton (i.e. 
Thalysias condition) etc., whereas in 5 species 
these differences are more subtle. 

Clathria (T) oxitoxa Levi (1963) is erect, 
bushy, flattened lamellate branches, yellow alive; 
skeleton plumo-reticulate \\ith fibres irregularly 
cored by very large principal styles and bundles 
(dragmata) of large oxeote toxas scattered 
throughout the sponge skeleton; ectosomal 
skeleton with plumose brushes of both ectosomal 
and subectosomal styles; principal styles 
regularly curx ed, with smooth non-t>'lote bases 
(450-800x35-40); subectosomal auxiliar\' sub- 
tylostyles with microspined bases (350-500 x 9); 
ectosomal auxiliary subtylostyles similar 
(100-150x4); echinating acanthostyles entirely 
spined with large spines (75-90 x 10); palmate 
isochelae in 2 size classes (6 and 13-14); aecolada 
toxas nearly oxeote, ranging from hair-like and 
faintly curved arms (170-250x0.5-1), thickly 
oxeote with straight arms (750-1300x4-7) and 
extremely long oxeotes with straight amis and 
only slight central curvature (2400-3000 x 
10-11); smaller wing-shaped toxas with large 
central curvature (35-40x0.5). This species is 
most similar to C. (M.) richmondi in having 
exceptionally large aecolada toxas in dragmata 
forming skeletal bands, but differs significantly 
in its live colouration, growth form, skeletal 
structure and the geometry and size of all spicules 
(in particular the huge upper size range of 
aecolada toxas). 

Clathria (M.) densa (Burton, 1 959) is massive, 
dense choanosomal skeleton with semi-pluinose 



ascending spongin fibres cored by principal 
styles, interconnected by few transverse fibres 
and spicule tracts; dense ectosomal skeleton of 
auxiliary^ subtvlostyles; principal styles with 
subtylote spined bases (175-298 X 18-35); sub- 
ectosomal auxiliaiy subt\'lost>'les with thick spined 
bases (130-275x4-8), echinating acanthost}4es 
thick, slightly curved, heavily spined with aspinose 
neck (11 8-1 56 X 9-16); palmate isochelae (9-13); 
hair-like aecolada toxas, distinctly sinuous and 
raphidiform forming dragmata (205-305 x 
0.5-1.5). This species is a borderline case 
between the subgenera Microciona and Clathria 
given that its choanosomal skeleton is a 
well-developed microcionid architecture that is 
verging on reticulate given the existence of 
vestigial inteconnecting fibres and spicule tracts. 
It also differs from C. (M.) richmondi in spicule 
geometT>', spicule sizes, and absence of smaller 
wing-shaped toxas: in fact the two species are 
only similar in possessing aecolada toxas 
forming dragmata. 

Clathria (C.) inhacensis Thomas (1979b) is 
thinly encrusting, surface conulose; ectosome 
reduced; choanosomal skeleton reticulate, with 
well developed ascending primary fibres cored 
by plumose tracts of principal styles, inter- 
connected by thinner transverse fibres in which 
few spicules are found and fiilly embedded with- 
in fibres, and both echinated by acanthostyles; 
subtylostyles interstitial and in brushes arising 
from tips of main fibres; principal styles with 
smooth bases (121-172x4-5); subectosomal 
subtylostyles with smooth bases (124-181 x 
2-4); echinating acanthostyles with variably 
spined shaft and spined bases (41-58x3-5); 
palmate isochelae (8-10); aecolada toxas hair- 
like (1 10-145 X 0.5-1 .5). This species clearly sits 
within subgenus Clathria given its possession of 
a reticulate skeletal architecture. It also ditfers 
significantly from C (M.) richmondi in spicule 
geometr>; spicule size and absence of toxodragmata. 

Clathria (T.) longitoxa (Hentschel, 1912) 
ranges from thinly encrusting to massive growth 
form; hymedesmioid to closely reticulate 
skeleton of stout fibres, with larger and smaller 
principal styles coring fibres in plumose arrange- 
ment, echinated by acanthostjdes; subectosomal 
auxiliary subtylostyles scattered; principal styles 
curved with smooth non-t>'lote bases (592-840 x 
22-26); smaller principal styles with subtylote 
granular bases (1 20-408 x 8-20); subectosomal 
auxiliary' subtylostyles with faintly microspined 
bases (430-584x4-9); ectosomal auxiliary 
subt>lostyles similar (190-320x3-5); echinating 



434 



MEMOIRS OF THE QUEENSLAND MUSEUM 



acanthostyles evenly spined 
with spined points (64-80 x 
6-7); palmate isochelae 
(12-20); accolada toxas with 
central U-bend and straight 
arms (400-820 x 1-2). This 
species differs from C. (M.) 
richmondi in most respects, 
showing similarities only in 
growth form, skeletal structure 
and possession of accolada 
toxas. 

Clathr'ta (C.) oculata 
Burton (1933a) has an erect 
branching anastomosing 
growth form, drab colouration 
with tinges of occasional 
purple; skeletal architecture 
composed of a subisodictyal 
reticulation of spongin fibres 
fully cored by principal styles 
and evenly echinated by 
acanthostyles; principal styles 
with smooth non-tylote bases 
(140-7); subectosomal auxil- 
iary subtylostyles with smooth 
bases (160x3); echinating 
acanthostyles evenly spined 
with small spines (65x4); 
accolada toxas slightly curved 
(160 long); palmate isochelae 
very small (6 long). This 
species is only similar to C 
(M.) richmondi in possessing 
accolada toxas, differing in 
most other features. 

In addition to these species 
there are three unnamed 
species described from Mada- 
gascar by Vacelet & Vasseur 
(1971) showing similarities to 
C. richmondi in the 
morphology of their accolada 
toxas, skeletal structure and 
growth form, although 
differing in most all other 
characters. 




FIG 4. FLto.somai bkclcion oi Claihria (Microciona) richmondi sp. nov. 
(holotype QMG306785). A, Bundles of subectosomal auxiliary subtylo- 
styles paratangential to and protruding through the surface, loosely 
associated with erect principal styles. B, Toxodragmata (bundles of 
accolada toxa) lying on or close to the surface. 



Clathria (T.) sp. 4 (Vacelet & Vasseur, 1971; 
see Table 2) is thinly encrusting, yellow alive; 
choanosomal skeleton microcionid with columns 
of fibres cored by principal styles and acantho- 
styles; ectosomal specialisation with some 
surface brushes but these are not thick; principal 
styles very slightly subtylote, smooth bases 



(130-440 X 6-12); subectosomal auxiliary subtylo- 
styles with smooth bases (150-320 x 4); ectosomal 
auxiliary subtylostyles with microspined bases 
(110-200x3); echinating acanthostyles slightly 
subtylote, poorly developed spines (55-60x5); 
palmate isochelae in 2 size classes, the smaller 
contort (5 and 12.5 long); accolada toxas nearly 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



435 



TABLE 2. List of Clathria species recorded from the Western Indian Ocean, Southeast Africa and Arabian 
Gulf-Red Sea provinces. Refer to Hooper (Hooper, 1996a) for full synonymy and taxonomic references. 
Annotation: 1 = species with accolada toxas; 2 ^ encrusting species with hymedesmioid-microcionid skeletal 
structure; 3 ^ identification has yet to be confirmed from examination of voucher specimen; 4 ^ identification 
unconfirmed, specimen voucher material missing; 5 = new combination; 6 = currently unrecognisable. 



Current taxonomic 
assignment 


Published name 


Author 


Western Indian Ocean 
records 


Other known distribution 


C. (Thalysias) abietina 
(Lamarck) 


C aculeata Ridley 


Burton (1959), 
Vacelet etal. (1976, 1977) 


Red Sea, S Arabian coast, 
Madagascar 


Tropical Australia, central 
NW Pacific, Philippines 


C. (Microciona) qffinis 
(Carter)^ 


M. affinis Carter 


Burton (1959) 


S Arabian coast, Zanzibar 


Gulf ofManaar 


C. (Thalysias) 
amirantiensis Hooper ' 


Colloclathria ramosa 
Dendy (preocc.) 


Dendy (1922), 
Hooper (1996) 


Amirante, Coetivy, 
Seychelles 




C. (Thalysias) anomah 
(Burton) ' 


R. anomala Burton 


Burton (1933) 


S South Africa 


- 


C. (Thalysias) anonyms 
(Burton) ^"^ 


M. anonyma Burton 


Burton (1959) 


Zanzibar 




C. (Clathf'ia) arbusculsi 
(Row) 


Ophlitaspongia arbusculsi 
Row, 0. horrida Row 


Row (1911) 


Red Sea 




C. (Microciona) 
atrasanguinea 
(Bowerbank) ^ 


M. atrasanguinea 
Bowerbank 


Carter (1880), 
Denrlv ('19'^?^ 

Burton & Rao (1932), 
Levi (1965), 
Van Soest(1993) 


Seychelles, Red Sea, Ara- 
bian Sea, Mauritius 


Caribbean, NE Atlantic, 
Mediterranean coast of 
India, Gulf of Manaar, 
Bay of Bengal, Andaman 
Sea 


C. (Clathria) axociom 


C. axociona Levi 


Levi (1963) 


S South Africa 


Namibia 


C. (Thalysias) cactiformis 
(Lamarck) ' 


Rhaphidophlus t\'picus 

(Carter), C. (T.) 
cactiformis (Lamarck), 
Rhaphidophlus sp. 2; 
Vacelet & Vasseur 


Vacelet etal. (1971, 1976, 
1 y / / ), riooper ( i yyo ) 


Madagascar, Somalia, E 
Africa, Seychelles, Red 
Sea 


S, W&Ecoasts ofAus- 


? C. (Clathria) caespes 
(Ehlers) " 


Scopalina caespes 
(Ehlers) 


Hooper (1996) 


S South Africa 


- 


C. (Wilsonella) 
cercidochela Vacelet & 
Vasseur 


Clathriopsamma 
cercidochela Vacelet & 
Vasseur 


Vacelet et al. (1971, 1977) 


Madagascar 


- 


C (Clathria) conica Levi 


C. conica Levi 


Levi (1963) 


S South Africa 


- 


C. (Thalysias) 
cuUingworthi Burton 


C cuUingworthi Burton 


Burton (1931) 


Natal 




C. (Clathria) dayi Levi 


C. dayi Levi 


Levi (1963) 


S South Africa 


(? Korea ^) 


C. (Thalysias) 
delaiibenfelsi (Levi) 


Rhaphidophlus 
delauhenfelsi Levi 


Levi (1963) 


S South Africa 




C. (Microciona) densa 
(Burton) ^"^ 


A/, densa Burton 


Burton (1959) 


S Arabian coast 




C. (Clathria) elastica Levi 


C. elastica Levi 


Levi (1963) 


S South Africa 


- 


C. (Axociella) fauroti 
(Topsent) 


Axosuberites fauroti 
Topsent 


Topsent (1893) 


Gulf of Aden 


- 


C. (Thalysias) flahellata 
(Burton) 


Rhaphidophlus flahellata 
Burton 


Burton (1936) 


S South Africa 




C. (Clathria) foliascens 
Vacelet & Vasseur 


C. foliascens Vacelet 
&Vasseur 


Vacelet etal. (1971, 1976, 
1977) 


Madagascar 




C (Thalysias) fusterna 
Hooper 


C. fusterna Hooper 


Hooper (1996) 


Eritrea 


N & NE Australia 


C. (Clathria) 
hexagonopora Levi 


C. hexagonopora Levi 


Levi (1963) 


S South Africa 




C. (Clathria) indica 
Dendy 


C. /«c//ca Dendy 


Burton (1931), Thomas 
(1979) 


Natal, Mozambique 


SE India, Gulf of Manaar 


C. (Clathria) inhacensis 
Thomas ' 


C inhacensis Thomas 


Thomas (1979) 


Mozambique 




C. (Clathria) irregularis 
(Burton) 


Marleyia irregularis 
Burton 


Burton (1931) 


Natal 





436 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 2. (cont) 



Current taxonomic 
assignment 


Published name 


Author 


Western Indian Ocean 
records 


Other icnown distribution 


C. (Clathria) Juncea 
Burton 


C juncsa Burton 


Burton ( 1 93 1 ) 


Natal 




C. (Microciona) 
laevissima (Dendy) ^ 


hi iaevissima Dendy 


Dendy (1922) 


]Vf auritius 




C (Thalysias) lambda, 
(Levi) 


Leptoclathria lambda 
Levi 


I evi (^1958^ 


Red Sea 




C. (Thalysias) lendenfeldi 
Ridley & Dendy ' 


C. spicata Hallmann, 
C. whiteleggii Dendy 


Dendy (1922), 
Burton (1931, 1959), 
Hooper (1996) 


Red Sea, Gulf of Aden, S 
Arabian coast, Cargados 
Carajos, Saya de Malha, 
Somalia, Natal 


SE, NE,N. &NW 
Australia, E Indonesia, 
Andaman Sea, Gulf of 
Manaar 


C, (Thalysias) lissoclada 
(Burton) 


Rhaphidophlus lissocladus 
Burton 


Levi (1963) 


S South Africa 


Falkland Is 


C. (Clathria) lobata 
Vosmaer 


C. lobata Vosmaer 


Vosmaer (1880). 
Ridley & Dendy (1887), 
Stephens (1915), 
Levi (1963) 


S South Africa 


- 


C. (Thalysias) longistyla 
(Burton) ^'^ 


M longistyla Burton 


Burton (1959) 


S Arabian coast 


(? Korea ^) 


C (Thalysias) longitoxa 
(Hentschel) 


M. longitoxa (Hentschel) 


Burton (1959) 


Gulf of Aden 


E Indonesia, Madras 


C. (Microciona) microxea 
(Vacelet & Vasseur) 2 


Paratenaciella microxea 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 




C. (Wiisonella) mixta 
Hentschel 


C mixta Hentschel 


Burton (1959) 


S Arabian coast 


E Indonesia ^ 


C. (Thalysias) nervosa 
(Levi) 


Axociella nervosa Levi 


Levi (1963) 


S Soudi Africa 


- 


C. (Clathria) oculata 
Burton ' 


C oculata Burton 


Burton (1933, 1959) 


Natal 


- 


C. (Thalysias) oxitoxa 


Cm oxitoxa Levi 




<1 ^mith Afrirn 




C. (Clathria) pachystyla 
Levi 


C pachystvla Levi 


Levi (1963) 


S South Africa 




C. (Axociella) parva Levi 


C. parva Levi 


Levi (1963) 


S South Africa 


Namibia 


C. (Thalysias) procera 
(Ridley) 


Rhaphidophlus procera 
Ridley, Echinonema 
gracilis Ridley 


i\iuicy yioot^, iviuicy oc 
Dendy (1887), Dendy 
(1922), Burton & Rao 
(1932), Burton (1931, 
1959), Levi (1963), 
Thomas (1973) 


Cargados Carajos, Sey- 
chelles, Amirante, Red 
Sea, Arabian coast, Natal 


Nh, N & iNW Australia, 
E Indonesia, Gulf of 
Manaar, (? Hawaii ^) 


C. (Clathria) rhaphidotoxa 
Stephens ' 


C. rhaphidotoxa Stephens 


Stephens (1915), 
Levi (1963) 


S South Africa 


- 


C. (Microciona) 
rhopalophora (Hentschel) ^ 


M. rhopalophora 
(Hentschel) 


Burton (1959) 


Maldives 


E. Indonesia, 
Cocos-Keeling, Gulf of 
Manaar 


C. (Thalysias) robusta 
(Dendy) 


M. robusta Dendy 


Dendy (1922) 


Amirante 


Singapore 


C. (Microciona) seriata 
(Grant) 


Ophlitaspongia seriata 
(Grant) 


Levi (1963) 


S South Africa 


NE. Atlantic, 
Mediterranean, New 
Zealand 


C. (Clathria) spongodes 
Dendy 


C. spongodes Dendy, 
C spongiosa Burton, 
C. madrepora Dendy 


Dendy (1922), 
Burton (1959), 
Vacelet et al. (1976) 


Red Sea, Gulf of Aden, 
Amirante, Madagascar, 
Seychelles 


(? Korea ^) 


C. (Microciona) 
stephensae Hooper 


M. similis Stephens 
(preocc.) 


Stephens (1915) 


S South Africa 




C. (Microciona) tenuis 
(Stephens) ^ 


M tenuis Stephens 


Stephens (1915) 


S South Africa 




C. (Clathria) transiens 
Hallmann ^ 


C. transiens Hallmann 


Burton (1959) 


Red Sea 


S Australian provinces 



A NEW CLA THRIA FROM THE INDIAN OCEAN 



437 



TABLE 2. (conO 



Current taxonomic 
assignment 


Published name 


Author 


Western Indian Ocean 
records 


Other known distribution 


C. (Clathria) typico. 
Kirkpatrick (virtually 
unrecognisable) 


C. typica Kirkpatrick 


Kirkpatrick (1904) 


Natal 




C (Microciona) vacehltia 
Hooper ^ 


A/, ciin'ichela Vacelet & 
Vasseur (preocc.) 


Vacelet & Vasseur 
(1965), Hooper(l996) 


Madagascar 


- 


C. (Thalysias) vulpi'na 
(Lamarck) ' 


C frondifera 
(Bowerbank), 
C. dichela (Hentschei) 


Ridley (1884), 
Ridley & Dendy ( 1 887), 
Topsent( 1892), Row 
(1911), Burton (1959), 
Levi (1961). Thomas 
(1973, 1979). Vacelet et 
al. (1971, 1976, 1977). 
rUUtzer-rmau yiyyj). 
Hooper ( 1 996), Kelly 
(1997) 


Madagascar, Aniirante, 
Seychelles, Red Sea, 
Mozambique, Aldabra, 
Zanzibar 


Tropical Australia, W & E 
coasts of India, Gulf of 

Manaar, Mergui 
Archipelago, Andaman 
Sea, Malaysia, h & W 
Indonesia, N Papua New 

Guinea, Vietnam, 
Philippines, Micronesia, 
S Japan, New Caledonia 


C. (Clathria) zoanthifera 
Levi 


C, zoanlhifera Levi 


Levi (1963) 


S South Africa 


- 


Clathria {Thalysias) sp.; 
Vacelet& Vasseur 


Rhaphidophlns sp. 1 ; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 


- 


Clathria (Thalysias) sp.; 
Vacelet & Vasseur 


Rhaphidophliis sp. 3; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 


- 


Clathria (Thalysias) s^.; 
Vacelet & Vasseur 


Rhaphidophliis sp. 4; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 




Clathria (Thalysias) s^.; 
Vacelet & Vasseur ' " 


Rhaphidophliis sp. 5; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 


- 


Clathria (Thalysias) s^.; 
Vacelet & Vasseur 


Rhaphidophlus sp. 6; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 


- 


Clathria (Tlialysias) sp.; 
Vacelet & Vasseur 


Rhaphidophliis sp. 7; 
Vacelet & Vasseur 


Vacelet etal. (1971, 1977) 


Madagascar 




Clathria (Microciona) sp.; 
Vacelet & Vasseur ^ 


Microciona sp. 1 ; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 




Clathria (Microciona) sp.; 
Vacelet & Vasseur 


Microciona sp. 2; 
Vacelet & Vasseur 


Vacelet & Vasseur (1971) 


Madagascar 




Clathria (Microciona) sp.; 
Vacelet & Vasseur " 


Microciona sp. 3; 
Vacelet & Vasseur 


Vacelet etal. (1971, 1976) 


Madagascar 




Clathria (Microciona) sp.^ 
Vacelet, Vasseur & Levi 


Microciona sp. 4; 
Vacelet, Vasseur & Levi 


Vacelet et al. (1976) 


Madagascar 





oxeote, with slight to virtually no central 
curvature (85-820x 0.5-2.5); U-shaped to 
wing-shaped toxas very small (5-7.5). 

Clathria (7:) sp. 5 (Vacelet & Vasseur, 1971) is 
encrusting, orange alive; hymedesmioid skeleton 
of principal subtylostyles and acanthostyles erect 
on basal spongin fibres; subectosomal and 
ectosomal auxiliary subtylostyles differ only in 
size and both contribute to both subectosomal 
and ectosomal surface brushes; principal 
subtylostyles slender, moderately subtylote micro- 
spined bases (140-280x5-8); subectosomal 
auxiliary subtylostyles slightly subtylote, smooth 
bases (up to 360x3); ectosomal auxiliary 
subtylostyles similar (from 90x2); echinating 
acanthostyles slender, evenly spined (50-60 x 
2-3); palmate isochelae in two size classes (4-5 
and 12-12.5 long); accolada toxas nearly oxeote, 



with straight or only very slightly curved arms 
and shght angular central flexion (35-250 long). 

Clathria (M ) sp. 2 (Vacelet & Vasseur, 1 97 1 ) is 
encrusting, pinkish to red alive, with white 
subectosomal drainage canals clearly visible on 
the otherwise smooth surface; skeleton micro- 
cionid; principal subtylostyles with smooth or 
microspined bases (330-550 x 13-15); subecto- 
somal auxiliary subtylostyles with smooth bases 
(120-550x2.5-5); echinating acanthostyles 
slender, entirely spined (10l)-l 20 x 5-7.5); 
palmate isochelae (7.5-20); accolada toxas with 
only slight curvature of arms and central flexion 
(130-320); small oxhom toxas (7.5-20). 

LITERATURE CITED 

AKNIN, M., GAYDOU,E.M., BOURYESNAULXN., 
COSTANTINO, V., FATTORUSSO, E. & 
MANGONI, A. 1996. Nor sterols in Axinella 



436 



MEMOUiS OK Tllb gUhbiNSLAND MUSLUM 



piiili tVi'ons, spongo f'nuii thr Iruii.in '. h L-an. 
Coiiipuiativc Hiochci)lisu> tmd i^hysioivc)' B 
HiocHemislry & Mo^l^eufar BiolQSy Ulbt4): 
845-848. 

BAER^ L. ]905.Sil}cispoDBicn von Sansibai^Kflpsmdt 

BARICAI. A.. OAVIS. r.L. ^ TtiGWH I.. S, 1996. 
Prey selecUDii by ihc Souili African Cape ruck 
lobster .Josks JaUvnJii'. theological iind physi- 
oFouical approaches. Bullclin of Marine Science 

smy. 1-S. 

SAVES IRELI.O, (.,. CM l AN'Id Vlli Tll, R., 
CERR.\N(). r. ^ SAR\. M Spicule 
dissolution in living; U'thvu (>nuiricnsi.\(\\\niQXSL: 
Demospoimuie* from a tropical c-ave, Bul|d&l o£ 
Marine Science 58(2): 598-601. 

$EER,& &ILAK M. 1#98. bisiWffleasuremcnis i)f 
^otosynthetlc irradiance respMiss^ Of two Red 
S^ftsp^iiges growing un<ier cm l|el^c(»iditiQt)s. 
Uarhte Biology Ul(4>: 6l3r«tt 

l>X Dilemwaoftcs a C. umisuaJ tftdob 

nIkaloifH irom n mixed collection of South 
Atficdfi .sponges. Journal of Natural I'litducls 
t)U5i: fW ^Ol, 
BOROJLVIC. R. 1%7. Sponi^ijiras d^/Uriquc du Sud 
(2K;ilcarc:i. ftansaclions of the Rsyal Socte^of' 

BOSR MJC'. \-. l^M 'i. Die rciractincllidcn, R«ise-' 
Osiiiffika A. Vucllzkow 3:23 1-251. 

BOWERBAMIC, J.S. 1862. On ihi; anatomy and plo si- 
pJuiSy ul'the Spongiadae. Part lU: On the generic 
-characters, thcsp^ific characters and the mediod 
itif OjXami^aiiioiV PbOose^caL T(:^actjDas ori^k* 

BRAEKI^, j^^ DADOED., DB QROOTE, S., 
fKRNAM)ESs J,B; & SOEST, -R.W.M. VAM 

1 90S New polykdidcs from the sponiie Plukort.',^ 
)t\x .luurrial ofNatural Products 6 1 (S ): 1 03S- 1 042. 
BSUl'li. \J. I '^76. Disciiis mvitac sp. no\. a ricv\ 
sponge asiotiatc from Kenya (Decupoda. 
Matantia. OisciadidaeK Cru&tai::^&ria 31(2): 

BURTON. M. 1^:6. DL-.scriptions of South .Mrican 
sponges collected In die Soudi African inanne 
Surve). Pan I. M\ xosportgWa and Astroteira- 
xonida. Unwn of South Africa. Fisheries and 
Marine Biological Slu vcv Report 4: 1-29. 
1929. Dc^Hrripiioii of Suuiti AiVican JipQQi^es* 
tolleciedin ihe Sniiih Afir^Tnannpsurvii^xPafi 
II The Lilhistidac, with a critical survey 6f ihfi 
dtfOTa^-lbmimgspongiii^. Ufiiofi of South Africa. 
Fisheries and Marine BtDlOgtcal Survi^ ifitm 
f7>: M2. 

1931. On« collection of marine sponges mt> .i!> 
Proni (hf. Natal coast. Annals of the N'atai 
Museum 6Cl);337-35S, 

I^*j3a. i-ouj" new marine sponges from Natal, 
Artnal* of the Nata^ ^4usei^m 7(2): 249-254. 



! *-)33b, Rt jH-rt 1711 .1 srii.ill .ul led ion of sponges from 
Still Bas; Soutli -Vxica, AlVl^^'s ^"^^ Maga:?ine of 
Natural MSfltery(HJ) 3l< 555-244. 

tQ36. Notes on spDngf?s iVom :Sou4 AiVica, with 
descriptions of new species. AnTiflJft antf 
.Ma^iOfiofhUrtu^Histf^ 17: 14]^147^ 

Otejf^ter I. I»orfrera."Soutli Aftlcarr Aniuiftt 
Life. Vol. 5 (Almqvisi Sc Wiksell: Swedenl 

1959. Sponecs. In Scicniihc Reports of llie John 
Miiria> r-Npo..riTion !Vrn-.i4, Vol. 10 (Bntil^ 
MubCLini (Natural Histors ): Londtm). 
BURTON. M. iic RAO. ll.s! h^o. Rcpoil oti llic 
shallow-wuler marine sportges in the collection of 
ilu' Indian Museum. Records of the Indiaji 

Museum ^M^^y. 299o.'^(». 
•CARMni->. S., td.BREVrstS. r . KASHMAN, Y, 
SiCl:LTON, B.W., WfU ih, A.H. !Sl Y0SII:K T. 
I900_ Dysidamide, a novel metabolite from ;i Ked 
Seaspoogei>t.v/t/ai he/ktcta. Australian Jounud 
of(:hemisrf%"43(U): I«KMS88. 
CAKM6tY,S. *JM4«MAN*V. I9iE0.Neviotme A, a 

pew triterpen^ from the lled&a sponge Stpho- 
novhaJina siphonella. Jcuftifll of Organs 
CheiniJfitr>*51(6>: 784-788. 
CARTER. H,J. 1S69. A descriptive acoouTlt of four 
Arubian and firtti&h, With 
general oIvvCtA .itions. .\nnal$ .^tftd fcb^SlZlIDSl t>f 
Namral llisiury (4l 4: 1-28, 

1871 . Dcscripliou anJ ilKistrations of a new Npecios 
of leihya, v\ ith nbscrv atlDns on the nomcticlaiurc 
of the Telhyadae. \iinal> and Magi^inft ot 
N;iiural History (4)8: ^J':)-lli5. 

l8Kii- Sponges liricludiiiu list of sponge*; dre^get} 
by the Birmingham Nattiral History and Micro- 
scopical Society. Falmouth Excursion^ 1879. 
Depth 15^20 IF^thoois). Midland Naturalist 

DAY, J.H. 1981. Thecstuarine fauna. Pp. 147 178. In 
Estuarine ecology with particulaj- retercncc to 
soudicni Africa. (Balkcma: Rotterdam i, 

01-T AK^; R. 1'"'46. Animaux dc Madai^ascar. Annalt's 
du Miisec (- olonia! de Marseille ()(4): 107-228. 

DLLSlVni. t. .. KASIIMAN, Y. & IXIfRASSI. C. 
1979. l:rgo>lLio.7.'-)( 1 1 ). 22-lelraen-";B-o| and it.s 
24E-Eth> l Homol<ig, Two ne^v marine sterols 
from the Red Sea &p*_-tn^e Bjentna foUi^. Helvetica 
Chnnica AcUi 62(6): 20^7-2045. 

DI ND^'. A. !'>! X Report on the calcareous sponges 
collected hv 1 IMS Sealnrk' in the Indian Ocean. In 
Reports of the Percv Sladcii I rusl Li.Kpcdttion lo 

Die [jQduio Qoesa in l905« \^limie S^Transactions 
efibe Linritian Society orLfjiidtan, Xaology l& 

1-29. 

1915. Report on the calcareous sponges collected 
by Mr James Horncll at Okhamuodal in 
Kaniawar in 1 905- 1 ^06. In Report to the 
Government ol Bartida on the Marine Zoology of 
Okhamandal- \biume 2. 17: 78-91. 

I'M 6a. Report on the iiexaLtineiiid ^punges 
(triaxonida) collected by HMS Seaiark in the 
Indian Ocean* la Reports of th& P^^:^ S}9dea 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



439 



Trust Expedition to the Indian Ocean in 1905, 
Volume 6. Transactions of the Linnean Societ>' of 
London. Zoology 17:211 -224. 

1916b. Report on the Homosclerophora and 
Astroletraxonida collected by HMS Sealark in 
the Indian Ocean. In Reports of the Percy Sladen 
Trust Expedition to the Indian Ocean in 1905, 
Volume 6. Transactions of the Linnean Society of 
London, Zoology 17: 225-271. 

1916c. Report on the non-calcareous sponges 
collected by Mr James Hornel! at Okhamandal in 
Kattiawar in 1905-1906. In Report lo the 
Government of Baroda on the Marine Zoology of 
Okhamandal. Volume 2. 17: 96-146. 

1922. Report on the Sigmatotetraxonida collected 
by HMS Sealark in the Indian Ocean. In Reports 
of the Percy Sladen Trust Expedition lo the 
Indian Ocean in 1 905, Volume 7. Transactions of 
the Linnean Societ>' of London, Zoology 18: 
1-164. 

DUNCAN, P.M. 1880. On a parasitic sponge of the 
order Calcarea. Journal of the Royal Microscopic 
Society 3:377-383. 

EHLERS, F. 1870. Die Esper'schen Spongien. 
(Zoologischen Sammlung der K. Universitat 
Erlangen. Programin zum Eintiitt in der Senat der 
Koniglichen Friedrich-Alexanders-Universitat in 
Erlangen: Erlangen). 

FINAMORE, E., MINALE, L. & ALI MOHAMMED, 
M. 1983. The sterols of the Somalian sponge 
Pleraphysialla (Pleraplysilla} piapyraceu. 
Rendiconlo dell'Accademia delle Scienze Fisiche 
e Malematiche (Serie IV) 50; 81-86. 

GEBREYESUS, T, YOSIEF, T, CARMELY, S. & 
KASHMAN, Y. 1988. Dysidamide, a novel 
hexachloro metabolite from a Red Sea sponge 
Dvsidea sp. Tetrahedron Letters 29(31): 
3863-3864. 

GRAY, J.E. 1 873. Natal sponges. Annals and Magazine 
of Natural History (4) 12: 264. 

GUO, Y, GAVAGNIN, M., MOLLO, E., TRIVEL- 
LONE, E., CIMINO, G., HAMDY, N.A., 
FAKHR, 1. & PANSINI, M. 1996. A new 
norsesterterpene peroxide from a Red Sea sponge. 
Naairal Product Letters 9(2): 105-112. 

GUO, Y, GAVAGNIN, M., MOLLO, E. & CIMTNO, G 
1997a. Hurghamides A D, new N acyl 2 
methylene (beta) alanine methyl esters from Red 
Sea Hippospcm^^'ia sp. Natural Product Letters 
10(2): 143-150.' 

GUO, Y, GAVAGNIN, M., MOLLO, E., CIMINO, G, 
HAMDY, N.A., FARHR, L & PANSINI, M. 
1997b. Hurghamides A D, new N acyl 2 
methylene (beta) alanine methyl esters from Red 
Sea Hippospon^ia sp. Natural Product Letters 
9(4): 281-288. ' 

HENTSCHEL, E. 1912. Kiesel- und Homschwamme 
der Aru und Kei-lnseln. Abhandlungen 
Senckenbergiana nalurforschende Gessellschafi 
(1912): 295-448. 



HOOPER, G.J., DAVIES-COLEMAN, M.T., 
KELLY-BORGES, M. & COETZEE, P.S. 1996. 
New^ alkaloids from a South Afiican Latrunculid 
sponge. Tetrahedron Letters 37: 7135-7138. 

HOOPER, J.N.A. 1 994. Coral reef sponges of the Sahul 
Shel f- a case for habitat preservation. Memoirs of 
the Queensland Museum 36( I): 93-106. 
1996a. Revision of Microcionidae (Porifera: 
Poecilosclerida: Demospongiae), with 
description of Australian species. Memoirs of the 
Queensland Museum 40: 1-626. 
1996b. A toxic Biemna from Madagascar 
(Demospongiae: Poecilosclerida). In Willenz, P. 
(ed.) Recent Advances in Sponge Biodiversity 
Inventory and Documentation, Bulletin de 
rinstitut Royal des Sciences Naturelles de 
Beigique (Supplement) 66: 123-134. 

HOOPER, J.N. A., KENNEDY, J. A., LIST- 
ARMITAGE, S.E.. COOK, S.D. & QUINN, R. 
1999. Biodiversity, species composition and 
distribution of marine sponges in northeast 
Australia. Pp. 263-274. In Hooper, J.N.A. (ed.) 
Proceedings of the 5th International Sponge 
Symposium. Brisbane, June-July 1998. Memoirs 
of the Queensland Museum 44: 263-274. 

HOOPER, J.N.A. & KRASOCHIN, V.B. 1989. 
Redescription of the burrowing sponge Zyzzya 
massalis (Dendy) from the Seychelles and 
Houtman-Abrolhos Islands. The Beagle, Records 
of the Northern Territory Museum of Arts and 
Sciences 6(1): 133-140. 

HOOPER, J.N.A. & LEVI, C. 1994. Biogeography of 
Indo-west Pacific sponges: Microcionidae, 
Raspailiidae, .Axinellidae. Pp. 191-212. In Soest, 
R.W.M. van., Kempen, TM.G van & Braekjnan, 
J.-C. (ed.) Sponges in Time and Space (Balkema: 
Rotterdam). 

HOWSON, CM. & CHAMBERS, S.J. 1999. Ophli- 
taspongia and Ophlitaspongia papilla reinstated, 
and a new species of Ophlitaspongia described 
(Porifera: Demospongiae: Microcionidae). 
Journal of the Marine Biological Association of 
the United Kingdom 79: 609-620. 

ILAN, M. & LOYA, Y. 1988. Reproduction and 
settlement of the coral reef sponge Niphates sp. 
(Red Sea). Pp. 745-749. In Choat, J.H., Barnes, D. 
& Borowitzka, M.A. (eds) Proceedings of the 
Sixth International Coral Reef Symposium. 
Townsville, 8-12 August 1988. ICRS, 6th 
International Coral Reef Symposium Executive 
Committee 2: 745-749. 
1 990. Sexual reproduction and settlement of the 
coral reef sponge Chalimda sp from the Red Sea, 
Marine Biologv 105: 25-31. 

ISAACS, S., BERMAN, R., KASHMAN, Y., 
GEBREYESUS, T & YOSIEF, T 1991. New 
polyhydroxy sterols, dysidamide, and a dideoxy- 
hexose from the sponge Dysidea herhacea. 
Journal of Natural Products, Lloydia 54: 83-91. 

1S.\.ACS, S. & K.ASHMAN, Y 1992. Shaagrockol B 
and C; two hexaprenylhydroquinone disulfates 



440 



MEMOIRS OF THE QUEENSLAND MUSEUM 



from the Red Sea sponge Toxiclona toxins. 
Tetrahedron Letters 33( 16): llll-ll'}^). 
IVANOVA, E.P., MIKHAILOV, V.V., KOUZNET- 
SOVA, T.A., AFIYATULLOV, A. A.. 
KALINOVSKAYA, N.I.. ELYAKOV, G.B., 
KIPRIANOVA, E.A. & GARAGULYA, A.D. 
1993. Heterotrophic bacteria associated with the 
sponge Dendrilla sp. and their physiological 
activity. Biologiya Morya, Vladivostok (1993): 
3-10. 

JAMES, D.M., KUNZE, H.B. & FAULKNER D.J. 
1991. Two new brominated tyrosine derivatives 
from the sponge Dnanella (=Psammaplysilla) 
purpurea. Journal of Natural Products. Lloydia 
54: 1137-1140. 

JAMES, P.S.B.R., GOPINADHA PILLAI, C.S., 
THOMAS, RA., JAMES, D.B. & KOYA, S. 
1989. 19. Environmental damage and con- 
sequences. Central Marine Fisheries Institute 
(1989): 212-227. 

JENKIN, C.F 1908. The calcareous sponges in the 
marine fauna of Zanzibar and British East Africa, 
from collections made by Cyril Crossland, M.A., 
in the years 1901 and 1902. Proceedings of the 
Zoological Society of London (1908): 434-456. 

KAMAT, S.Y., SOLIMABI, NAQVI, S.W.A., 
FERNANDES, L., REDDY, C.V.G, BHAKUNl, 
D.S. & DHAWAN, B.N. 1981. Bioactivity of the 
extracts from some marine animals of the Indian 
coast. Mahasagar 14(2): 117-122. 

KASHMAN, Y., GROWEISS, A., CARMELY, S., 
KINAMONl, Z., CZARKIE, D. & ROTEM, M. 
1 982. Recent research in marine natural products 
from the Red Sea. Pure and Applied Chemistry 
54(10): 1995-2010. 

KASHMAN, Y., CARMELY, S.. BLASBERGER, D., 
HIRSCH, S. & GREEN, D. 1989. Marine natural 
products: new results from Red Sea invertebrates. 
Pure and Applied Chemistry 61(3): 517-520. 

KELLER, C. 1889- Die Spongienfauna des rothen 
Meeres. 1 Halfte. Zeitschrift fur Wissenschaflliche 
Zoologie 48: 311-406. 
1891. Die Spongienfauna des rothen Meeres. 11 
Halfte. Zeitschrift fur Wissenschaflliche 
Zoologie 52:294-368. 

KELLY, M. 1997. Porifera. Sponges. Pp. 106-117. In 
Richmond, M. D. (ed.) A guide to the seashores of 
Eastern Africa and the Western Indian Ocean 
islands (Sida, Department for Research Co- 
operation, SAREC. Ord & Vetande AB: Sweden). 

KELLY-BORGES, M. & VACELET, J. 1995. A 
revision of Diacarnus Burton and Negomhata de 
Laubenfels (Demospongiac: Latrunculiidae) with 
descriptions of new species from the west central 
Pacific and the Red Sea. Memoirs of the 
Queensland Museum 38(2): 477-503. 

KIRKP.AfRICK. R. 1900. Note on the occurrence of the 
euplecteliid sponge Regadrella phoenix O. 
Schmidt, ofF the South African coast. Annals of 
the South African Museum 13: 63-64. 



1901. Description of a new hexactinellid sponge 
from South Africa. Annals and Magazine of 
Natural History (7) 7: 457-459. 

19()2a. Descriptions of South African sponges. 
Cape of Good Hope, Department of Agricuhure 
Bulletin. Marine Investigations in South Africa 
2(14): 219-232. 

1902b. Descriptions of South African sponges. Part 

II. Cape of Good Hope, Department of 
Agriculture Bulletin. Marine Investigations in 
South Africa 2(1 4): 171-180. 

1903. Descriptions of South African sponges. Part 

III. Cape of Good Hope, Department of 
Agriculture Bulletin. Marine Investigations in 
South Africa 2(1 6): 233-264. 

1904. Sponges. In Zoological Record (for 1902). 
Abstract by R. von Lendenfeld. Zoologische 
Zentralblatt 10: 147-148. 

1908. Description of a new dictyonine sponge from 
the Indian Ocean. Records of the Indian Museum 
2: 21-26. 

1913. Note on the occurrence of the euplecteliid 
sponge Regadrella phoenix, O. Schmidt, off the 
South African coast. Annals of the South Africa 
Museum 13: 64-66. 
KOBAYASHI, M., CHAVAKULA, R., MURATA, O. 
& SARMA, N.S. 1992a. Marine terpenes and 
terpenoids. XIV. Absolute configuration and acid 
catalyzed transfomiation of (-)- 1 2, 1 3-didehydro- 
furospongin-1 isolated from an Arabian sea 
sponge, Fasciospongia cavernosa Schmidt. 
Chemical Pharmaceutical Bulletin (Tokyo) 40: 
599-601. 

KOBAYASHI, M., KRISHNA, M.M., ISHIDA, K. & 
ANJANEYULU, V. 1992b. Marine sterols. 22. 
Occurrence of 3 oxo 4,6,8(14) tri unsaturated 
steroids in the sponge Dysidea herbacea. 
Chemical Pharmaceutical Bulletin (Tokyo) 40: 
72-74. 

KOLBASOV, G.A. 1990. Acasta pertusa sp. n. 
(Cirripedia. Thoracica) from the Red Sea. 
Zoologicheskii Zhumal 69(9): 142-145. 
1992. Two new species of the genus Acasta 
(Cirripedia, Thoracica) from the south western 
part of the Indian Ocean. Zoologicheskii Zhurnal 
71(1): 140-145. 

KONDRACKI, M.L. & GUYOT, M. 1987. 
Smenospongine: a c\iotoxic and antimicrobial 
aminoquinone isolated from Smenospongia sp. 
Tetrahedron Letters 28(47): 58 1 5-58 1 8. 

KOREN GOLDSHLAGER, G, KASHMAN, Y. & 
SCHLEYER, M. 1998. Haliclorensin, a novel 
diamino alkaloid from the marine sponge 
Haliclona tulearensis. Journal of Natural 
Products 61(2): 282-284. 

KUMAR, A. 1924a. A new variet>' of Leucosolenia 
gardineri (Dendy), Leiicosolenia gardineri van 
vergensis. Proceedings of the Lahore Philo- 
sophical Society 3: 21-22. 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



441 



1924b. On a probable new genus of marine sponge 
from Karachi. Proceedings of the Indian Science 
Congress 10: 1-167. 

1924c. Porifera from Karachi. Proceedings of the 
Lahore Philosophical Societ>^ 3: 67-68. 

1925. Report on some tetraxonid sponges in the 
collection of the Indian Museum. Records of the 
Indian Museum 27: 21 1-227. 
LAGHI, GF., MARTINELLI. G & RUSSO, F 1984. 
Localization of minor elements by EDS 
microanalysis in aragonilic sponges from the St 
Cassian Beds, Italian Dolomites. Lethaia 17(2): 
133-138- 

LENDENFELD, R.VON 1 897. Spongien von Sansibar. 
Abhandlungen Senckenbergiana 
Naturforschende Gessellshcaft 21 : 93-133. 

LEVI, C. 1956. Spongiaires des cotes de Madagascar. 
Memoires de Tlnslitut Scientifique de Mada- 
gascar (A) 10: 1-23. 
1958. Resultats scientifiques des campagnes de la 
Calypso'. Fascicule III. V. Campagne 195 1-1952 
en Mer Rouge. 5. Spongiaires de Mer Rouge 
recueillis par la Calypso' (1951-1952). Annates 
del'Institut Oceanographiquc, Monaco 34: 3-46. 
1961. Resultats scientifique des campagnes de la 
Calypso'. Fascicule V. XIV. Campagne 1954 
dans rOcean Indien. 2. Les spongiaires de Pile 
Aldabra Campaigne Oceanographiquc de la 
Calypso' (May-Juin 1954). Annales de I'lnstitut 
Oceanographiquc, Monaco 39: 1-3L 

1963. Spongiaires d'Afrique du Sud (I) Poecilo- 
sclerides. Transactions of the Royal Society of 
South Africa 37(1): 1-72. 

1964. Spongiaires du canal de Mozambique. 
Bulletin du Museum National d'Histoire 
Naturelle 36(3): 384-395. 

1965. Spongiaires recolles par I'Expedition 
Isracliennc dans !e sudde la Mer Rouge en 1962. 
Bulletin of the Sea Fisheries Research Station, 
Israel. Israel South Red Sea Expedition, 1962, 
Report 13): 3-27. 

1967. Spongiaires d'Afrique du Sud (3) Tetractin- 
ellides. Transactions of the Roval Society of 
South Africa 37(3): 227-256. 

1986. Laocaetis perion nov. sp., spongiaire 
hexactinellide Craticulariidae de Focean Indien. 
Bulletin du Museum National d'Histoire 
Naturelle, Paris (4) 8(A, 3): 437-442. 
MAGNINO, G. & GAINO. E. 1998. HaplosyUis 
spongicola (Grube) (Polychaeta, Syllidae) 
associated with two species of sponges from East 
Africa (Tanzania, Indian Ocean). Marine Ecology 
19(2): 77-87. 

MARSDEN. J.R. 1975. Classes of lipids in marine 
sponges from Kenya. Journal of Experimental 
Marine Biology and Ecology 1 9: 9-18. 

McCABE, T.. CLARDY, J., MINALE, L., PIZZA, C, 
ZOLLO, F. & RICCIO, R. 1982. A triterpenoid 
pigment with the isomalabaricane skeleton from 
the marine sponge Stelletta sp. Tetrahedron 
Letters 23(33): 3307-3310. 



McPHAlL, K., DAVIES COLEMAN, M.T. & 
COETZEE, P. 1998. A new fliranosesterlerpene 
from the South African nudibranch Hypselodoris 
capensis and a Dictvoceratida sponge. Journal of 
Natural Products 6f(7): 961-964. 

MEBS, D. 1985. Chemical defense of a dorid 
nudibranch, Glossodoris qiiadricolor from the 
Red Sea. Journal of Chemical Ecology 11(6): 
713-716. 

MERGNER, H. 1982. Man made influences on and 
natural changes in the settlement of the Aqaba 
reefs (Red Sea). Proceedings of the International 
Coral Reef Symposium (4)T 193-207. 

NGOC HO, N. 1990. Nine Indo Pacific species of 
Upogebia Leach (Crustacea: Thalassinidea: 
Upogebiidae). Journal of Natural Histor>' 24(4): 
965-985. 

PARAMESWARAN, RS., DAS, B. & KAMAT, S.Y. 
1994. Lipid contents of the sponge Haliclona sp. 
Indian Journal of Chemistry (B) Organic 
Chemistry, Medicinal Chemistry' 33B(1 ): 99-101 . 

PARA MESWA RAN. P.S., KAMAT, S.Y., 
CH.ANDRAMOHAN. D., NAIR, S. & DAS, B. 
1992a. Anti-bacterial compounds from the 
sponge Haliclona sp. Pp. 4 1 7-420. In Desai, B. N. 
(ed.) Oceanography of the Indian Ocean (Oxford 
& IBH Publishing Co.: New Delhi, Bombay). 

PARAMESWARAN, P.S., NAIK, C.G, DAS, B. & 
KAMAT, S.Y. 1989. Sterols from the 
Lakshadweep sponge, Ircinia ramosa (Keller). 
Indian Journal of Chemistry Section (B) Organic 
Chemistry, Medicinal Chemistry 28(12): 1091- 
1092. 

1992b. Minor sterols from the sponge Ircinia 
ramosa (Keller). Pp. 413-416. In Desai, B. N. 
(ed.) Oceanography of the Indian Ocean (Oxford 
& IBH Publishing'Co.: New Delhi, Bombay). 

PARAMESWARAN, RS., NAIK, C.G. & ilEGDE, 
V.R. 1997. Secondary metabolites from the 
sponge Tedania anhelans: isolation and 
characterization of two novel pyrazole acids and 
other metabolites. Journal of Natural Products. 
Lloydia60(8): 802-803. 

PATEL, B., PATEL, S. & BALANl, M.C. 1985. Can a 
sponge fractionate isotopes? Proceedings of the 
Royal Society of London (B) Biological Sciences 
224(1234): 23-41. 

PETTIBONE, M.H. 1993. Scaled polychaeles 
(Poiynoidae) associated with ophiuroids and other 
invertebrates and review of species referred to 
Malmgrenia Mcintosh and replaced by 
Malmgreriiella \ lartman, with descriptions of new 
taxa. Smithsonian Contributions to Zoology 
(538): 1-92. 

PETTIT, G.R., CICHACZ, Z.A., RUI, T, HOARD, 
M.S., MELODY, N. & PETTIT, R.K. 1998a. 
Antineoplastic agents. 386. Isolation of 
sesterstalins 1 3 from the marine sponge Hyrtios 
erecra. Journal of Natural Products 61(1): 13-16. 

PETTIT, GR., GAO, F., DOUBEK, D.L., BOYD, 
M.R., HAMEL, E., BAL R.L., SCHMIDT, J.M., 



442 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TACKETT, L.P. & RUTZLER, K. 1993a. 
Antineoplastic agents. 252. Isolation and structure 
orHalistatin-2 from the Comoros Marine Sponge 
Axinella carteri. Gazzeti Chlmica Italiana 123: 
371-377. 

PETTIT, G.R., HERALD, C.L., CICHACZ, Z.A., 
GAO, F., SCHMIDT, J.M., BOYD, M.R., 
CHRISTIE, N.D. & BOETTNER, RE. 1993b. 
Isolation & structure of the powerful human 
cancer cell growth inhibitors spongistatins 4 & 5 
fi"om an African Spirustrella spimspiruUfera 
(Fori (era). Journal of the Chemical Society, 
Chemical Communications 1993: 1805-1807. 

PETTIT G.R., GAO, R, SCHMIDT J.M., CHAPUIS, 
J.-C. & CERNY, R.L. 1994a. Isolation and 
structure of Axinastatin 5 fi"om a Republic of 
Comoros marine sponge. Bioorganic and 
Medicinal Chemistry Letters 4(24): 2935-2940. 

PETTIT GR.. RUI, T, HERALD. D.L., CERNY, R.L. 
& WILLIAMS, M.D. I994h. Antineoplastic 
agents. 277. Isolation and structure of 
phakellistatin 3 and isophakellistatin 3 from a 
Republic of Comoros marine sponge, .fournal of 
Organic Chemistrv' 59(7): 1593-1595. 

PETTIT G.R., McNULTY, J., HERALD, D.L., 
DOUBEK, D.L., CHAPUIS, J.C., SCHMIDT, 
J.M., TACKETT, L.R & BOYD. M.R. 1997. 
Antineoplastic agents. 362. Isolation and X ray 
crystal structure of dibromophakcllstatin from the 
Indian Ocean sponge Phakellia mauritiana. 
Journal of Natural Products, Lloydia 60(2): 
180-183- 

PETTIT GR., TAN, R., MELODY, N., CICHACZ. 
Z.A., HERALD, D.L., HOARD, M.S., PETTIT 
R.K. & CHAPUIS, .I.e. 1998b. Antineoplastic 
agents. 397. Isolation and stiTJCture of sesterstatins 
4 and 5 from Hyrtios erecfa (The Republic of 
Maldives). Bioorganic Medical and Chemical 
Letters 8: 2093-2098. 

PULITZER-FINALI, G 1993. A collection of marine 
sponges from East Africa. Annali del Museo 
Civico di Storia Naturale G. Doria' 89: 247-350. 

RAHIM, M.A. 1979. Porifcra. Publications Saudi 
Biological Society 3: 46. 

RAMADAN, S.A. 1997. Two new species of meso- 
stigmatid mites (Acari) associated with sponges 
from the Red Sea, Egypt. Assiut Veterinary 
Medical Journal 38(75): 191-204. 

RICHMOND, M.D. 1997. A guide to the seashores of 
Eastern Africa and the Western Indian Ocean 
islands. (Sida, Department for Research Co- 
operation, SAJIEC. Ord & Vetande AB: Sweden). 

RIDLEY, S.O. 1884. Spongiida. Pp. 366-482, 582-635. 
In Report on the Zoological Collections made in 
the Indo-Pacific Ocean during the Voyage of 
H.M.S. Alert 1881-2 (British Museum (Natural 
History): London). 

RIDLEY, S.O. & DENDY, A. 1887. Report on the 
Monaxonida collected by HMS Challenger' 
during the Years 1 873-76. Pp. 1-275. In Report on 
the Scientific Results of the Voyage of H.M.S- 



Challenger during the Years 1873-76. 20 (Her 
Majesty's Stationery Office: London, Edinburgh, 
Dublin). 

RINKEVICH, B.. SHASHAR, N. & LIBERMAN, T. 
1993. Nontransitive xenogeneic interactions 
between four common Red Sea sessile 
invertebrates. Pp. 833-839. In Richmond, R.H. 
Proceedings of the Seventh International Coral 
Reef Symposium. Mangilao, Guam, 22-26 June 
1992. University of Guam Press 2: 833-839. 

ROW, R.W.H. 1911. Reports on the marine biolog\' of 
the Sudanese Red Sea, from collections made by 
Cyril Crossland, MA, BSc, FZS. XIX. Report on 
the sponges collected by Mr Cyril Crossland in 
1904-5. Part II. Non-Calcarea. Journal of the 
Linnean Societv, Zoology 3 1(208): 287-400. 

RUDI, A. & KASHMAN, Y. 1993. Aaptosine a new 
cytotoxic 5,8 diazabenz(cd)azulene alkaloid from 
the Red Sea sponge Aaptos aaptos. Tetrahedron 
Letters 34(29): 4683-4684. 

RUDI, A., TALPIR, R., KASHMAN, Y., BENAYAHU, 
Y. & SCHLEYER, M. 1993. Four new C16 1,2 
dioxene polyketides from the sponge Plakortis 
aff. .simplex. Journal of Natural Products, Lloydia 
56(12); 2178-2182. 

RUDI, A., KASHMAN, Y., BENAYAHU, Y. & 
SCHLEYER, M. 1994a. Sodwanones A F, new 
triterpenoids from the marine sponge Axinella 
weltmri. Journal of Natural Products, Lloydia 
57(10): 1416-1423. 

RUDI, A., STEIN, Z., GREEN, S., GOLDBERQ I., 
KASHMAN. Y., BENAYAHU, Y. & 
SCHLEYER, M. 1994b. Phorbazoles A D, novel 
chlorinated phenylpyrrolyloxazoles from the 
marine sponge Phorbas aff. clathrata. 
Tetrahedron Letters 35(16): 2448, 2589-2592. 

RUDI, A.. KASHMAN, Y, BENAYAHU. Y & 
SCHLEYER, M. 1995. Durbinal A, B and C: 
three new cytotoxic sponge metabolites. 
Tetrahedron Letters 36(27): 4853-4856. 

SAMAAI, T. 1997. Syslematics, phylogeny and 
biogeography of a selection of poecilosclerid 
sponges from Ouderkraal, on the west coast of 
South Africa. Unpubl. MSc thesis, Department of 
Biology, Imperial College, London. 

SARA, M. & BAVESTRELLO, G. 1995. Tethya 
omanensis, a remarkable new species from an 
Oman cave (Porifera, Demosponsiae). Bollettino 
diZoologia 62(1): 23-27. 

SAR.\, M. & CORRIERO, G 1994. A new species of 
Teiliya, T. temdsclera (Porifera Demospongiae) 
from the Maldives. Bollettino dei Musei e degli 
Istituli Biologici della Universita di Genova 
58-59: 69-75. 

SARA, M., CORRIERO, G & BAVESTRELLO, G 
1993a. Tethya (Porifera, Demospongiae) species 
coexisting in a Maldivian coral reef lagoon: 
taxonomical, genetic and ecological data. Marine 
Ecology 14(4): 341-355. 
1993b. Tethya peracnta (YopsQwi) SiX\d T comorensis 
n sp (Porifera, Demospongiae) from the coral 



A NEW CLATHRIA FROM THE INDIAN OCEAN 



443 



reef of Mayotle (Comoresj. Bollettino di 
Zoologia 60: 219-224. 

SARA, M., PANSINI, M. & PRONZATO, R. 1979. 
Zonation of photophilous sponges related to water 
movement in reef biotypes of Obhor Creek (Red 
Sea). In Levi, C. & Boury-Esnault, N. (eds) 
Biologic des Spongiaires. Sponge Biology. 
Colloques Intemalionaux du Centre National de la 
Recherche Scientifique (291): 282-288. 

SCHLEYER, M.H. 1991. Shell borers in the oyster, 
Striostreo margaritacea: pests or symbionts? 
Symbiosis 10(1-3): 135-144. 

SCHMIDT, E.O. 1862. Die Spongien des Adriatischen 
Meeres (Wilhelm Engelmann: Leipzig). 

SCHMIDT E.W., HARPER, M.K. & FAULKNER, 
D.J. 1997. Mozamides A and B, cyclic peptides 
from a theonellid sponge from Mozambique. 
Journal of Natural Products, Lloydia 60(8): 
779-782. 

SOEST R.W.M. van 1984. Marine sponges from 
Curasao and other Cai'ibbean localities. Pai1 III. 
Poecilosclerida. Studies on the Fauna of Curasao 
and other Caribbean Islands (199): 1-167. 
1993. Distribution of sponges on the Mauritanian 
continental shelf. In Wolff, W.J., van der Land, J., 
Nienhuis, RH. & de Wilde, P.A.W.J. (eds) 
Ecological studies in the coastal waters of 
Mauritania. Hydrobiologia: 258: 95-106. 
1994a. Demosponge distribution patterns. Pp. 
213-224. In Soest, R.W.M. van, Kempen, TM.G. 
van & Braekman, J.-C, (eds) Sponges in time and 
space. (Balkema: Rotterdam). 
1 994b. Sponges of the Seychelles. Pp. 65-74. In vain 
der Land. J. (ed.) Oceanic reefs of the Seychelles: 
report on a cruise of RV Tyro to the Seychelles in 
1992 and 1993 (National Museum of Natural 
History: Leiden). 

SOEST, R.W.M. VAN & HAJDU, E. 1 997. Maiine area 
relationships from twenty sponge phylogenies. A 
comparison of methods and coding strategies. 
Cladistics 13: 1-20. 

SOEST, R.W.M. VAN, ZEA, S. & KIELMAN, M. 
1994. New species of Zyzzya, Cornulella and 
Damiria (Porifera: Poecilosclerida), with a 
review of fistular genera of lophonidae. Bijdragen 
totde Dierkunde64(3): 163-192. 

SOKOLOFF, S., HALEVY, S., USIELI, V.. 
COLORNI, A. & SAREL, S. 1982. Prianicin A 
and B, nor seslerterpenoid pero.xide antibiotics 
from Red Sea sponges. Experientia, Basel 38: 
337-338, 

SOLLAS, I.B.J. 1908. The inclusion of foreign bodies 
by sponges, with a description of a new genus and 
species of Monaxonida. Annals and Magazine of 
Natural Histoiy (8) 1 : 395-40 1 . 

STEPHENS, J. 1915. Atlantic sponges collected by the 
Scottish National Antarctic Expedition. 
Transactions of the Royal Society of Edinburgh 
50: 423-467. 

THOMAS, RA. 1973. Marine Demospongiae of Mahe 
Island in the Seychelles Bank (Indian Ocean). 



Annales du Musee Royal de LAfrique 
Central-Tervulen, Belgique (8, Sciences 
Zooiogiques) (203): 1-91. 

1975. Boring sponges of Zuari and Mandovi 
Estuaries. Bulletin of the Department of Marine 
Science University of Cochin 7(1): 117-126. 

1976a. Marine Demospongiae of Zanzibar Island. 
Journal of the Marine Biologv Association of 
India 18(3): 448-460. 

1976b. The history of spongology of the Indian 
Ocean. Journal of the Marine Biological 
Association of India 18(3): 610-625 

1979a. Demospongiae of Minicoy Island (Indian 
Ocean) Part 1 - orders Keratosida and 
Haplosclerida. Journal of the Marine Biological 
Association of India 21(1-2): 10-16. 

1979b. Endectyon lamellosa n. sp., 
(Demospongiae: Poecilosclerida, Raspailiidae) 
from the Indian seas and a revised key to the 
Indian species Endectyon Topsent. Journal of 
the Marine Biological Association of India 
18(1): 169-172. 

1979c. Studies of sponges of Mozambique Channel 
1. Sponges of Inhaca Island. 2. Sponges of 
Mambone and Paradise Islands. Annales du 
Musee Royal de I'Afrique Centrale, Tervuren, 
Belgique (8, Sciences Zooiogiques). Koninklijk 
Museum Voor Midden-Afrika Tervuren Belgie 
Annalen Zoologische Wetenschappen (227): 
1-73. 

i980a. Demospongiae of Minicoy Island (Indian 
Ocean) Part 2 - order Poecilosclerida. Journal of 
the Marine Biological Association of India 
22(1-2): 1-7. 

1 980b. Demospongiae of Minicoy Island (Indian 
Ocean) Part 3 - orders Halichondrida, 
Hadromerida, Epipolasida and Choristida. 
Journal of the Vlarine Biological Association of 
India 22(1 -2): 8-20. 

1981a. Marine Demospongiae of Ras Iwatine 
(Kenya). Journal of the Vlarine Biological 
Association of India 18(3): 642-649. 

1981b. A second collection of marine Demo- 
spongiae from Mahe Island in the Seychelles 
Baiik (Indian Ocean). Koninklijk Museum Voor 
Midden-Afrika Tervuren Belgie Annalen 
Zoologische Wetenschappen (233): 1-54. 

1988. Sponge generated bioerosion in Lakshad- 
weep. Indian Council of Agricultural Research 
Marine Fisheries Information Service Technical 
and Extension Series (86): 20-26. 

1989. 13. Sponge fauna of Lakshadweep. Central 
Marine Fisheries Research Institute (1989): 
150-161. 

THOMAS, RA., GOPINADHA PILLAI, C.S. & 
RAJAGOPALAN, M.S. 1997. Demospongiae of 
the Gulf of Kutch. Journal of the Marine Biolog- 
ical Association of India 38(1-2): 124-132. 

THOMSON, W. 1 868. On the vitreous' sponges. Annals 
and Magazine ofNatural History (4) 1 : 1 14-132. 



444 



MEMOIRS OV THE QUEENSLAND MUSEUM 



tOpSENT, E. 1890. Etudes de Spongiaires. IF. 
Description d'une Lithistide molle 1^ HeUillon 
{Kalicipsis pemwllis nov. sp.). Revuedfej^Qji^e 
du Nord France 2(8): 294-296. 
1892. Eponges de la Mer Rouge* Memoiies dfi la 

SoQi^t^Zookigique de Franc$ 1:21-^^: 
IS93a.MisSk)nJSc56hlirique delVt43h. AlNflirf-a*i?t 
ties S^cNIf^ (Mqis-MeL r892i), Spbti^ialrG^. 
BuU^fin la Soi^t^ 2^6ldg1(!|Ue a& France 

lS93b. Kcfte stir quelqiies Eponges du Golfe de 
Tadjoura recufillics par M Ic Dr T,. Faurot. 
BullcTin de Societe zoologique de France 18: 

J RIMliRn;Lll. a & FAULKNER, DJ. I994. Six 
new diterpene isonitriles from the sponge 
Acanrhella cavernosa. Journal of Natural 
Products, I.Ioydia57(4): 501-506. 
VACtLFT. J. 1967a. Descriptions d"eponges 
Pharetronides aciucllcs dcs tunnels obscurs 
soLLs-recifaux de Tulear (Madagascar). Recueil 
des Tiavau-K de la Station MaFrrie d'Endourae- 
Marseille (6): 37-62. 

1967b. Qiielques eponges Pharelronides et 
Silico-Calcaires de groties sous-marines 
obscures. Recueil des Travaux de fa Statran- 
Marine d'Eiwioujorie 5S: 12 M32. 

1977. Eponges Pliffl^nides actuelles et sd^Jao- 

VACELET, J-! TIERCELRM, JJ. & GASSF, F. 199K 
The sponge DosiUa brouni (Spongi I lidae ) in Lak6 
Biuingo. Gregoiy Rift, K^ya. Hydtobielogia 

211(1): il-iS. 
VACELE T, J. & VASSCIJR, P 1^65. Spongiaires dfiS'- 
grolles el surplombs dcs rccifs de Tulwar 
(Madagasciirj. Recueil dcs Travaux de la Station 
Marine crbiidounie-Marscillc, Supplement (4): 
71-123. 

1966. Les tunnels ohsurs soua-recilaux dc Tulear 



378. In Abstracts of the Second Interriational 
Qcpanographlc Congress^ (Moscow). 

iWl. Sponges des T6cxfs cOraHiens de Tulwar 

(Madagascar). Tcthys, Supplement 1: 51-126. 

1977. Sponge distribtilion in coral reels and related 
areas in the vicinit> ofTulcar (Madagascar). Pp. 
113-117. In Proceedings of the 3rd (nlernational 
Coral Reef Symposium. Miami, Florida.. 
(Rosepsti^l School of M^rin^ atifi Atmosph^c 

VAtEigt; I, VASSI^R. P. A TEVi, C, 1976. 
Spongialre^ de la penie exieme dcs rccits 
coralliens de Tulear (sud-Ouesl de Madiigascar). 
Mcmoires du Museum National dMli.stoire 
Naturelle. Paris ( A, /oologie) 49: 1-116. 

VFKKATnSW.AJlLU, V., FAULKNER, DJ„ RIOS 
STFINFR, J.L.. CORCORAN, E. &CtAJtt)Y J. 
1 99 1 . SmcnochromBnps> unusual JoiSicfflcycIic 
sesquiterpene %^qUtaOlie4ertV£l[&Ate? friii a 
S&ycheUes sporige of the g^u^ St^^mpium^^ 
Joitaafor<)rganfcCheniWS6?^)r62? 

VHRON, J.F.N. 1986. Corals of Australia and the Indo- 
Pacific. (/Vngus 6i Robertson: Sydne>, London ). 

1993, A biogeographic database of hermaiypic 
corals. Species of the Central Indo-Pacific 
Genera of the world. Australian Institute of 
Marine Science Monograph Scries 10: 1-433, 

VINE, P. I9S6. Red Sea invertebrates. lIMMFL 
Publishing; London). 

VOSMAFR, GCJ. 1 880. The sponges of the Leyden 
Museum. 1. The family of the Desmacidinae. 
Notes iTpni ^ Leaden, Museum 2: 99-J 64. 

'WQRHgiIiq5> G l^ftff. ^he tfi&f tJave dwelling 
vltr?iCO|isetvative coralline demosponge 
AslrOscleta wflleyana Lister 1900 from the 

Indo-Pacillc. Facies3S: 1-88, 

WRIGHT, E.P. 1881. On a new genus and species of 
sponge (Alemo seychellensis) with supposed 
he^ogiorohi^ ^i4§. Transactions of the Irish 



NEWSI^CIES OF TOXIC TEDANIA FROM NORTHERN VANUATU (POfelFKRAt: 
DEMOSmNOIAlJ: POECiLOSCLERIDA: TBDANUDAE) 



fOHN A. K©«NfiDY ANO )OKN N;A. HQOPISR 



KcnnccK. J. A ^ Hoopei',{J.M,A.20ii>0 Cf6 3' ^p.xie:> ofto\ic T^iSwin Irdm Northern 
Vanuatu (Pontcra: Demo^^ngiatiu* PneotjoscLerida: Tedaniidae). Met^ir^ cftffe Queens-- 

IbdaniafTedaviajsirorigyloKfyla^^. nov, is described, compaied with 7I^W£f, Emd|hertOxif^ 
species &om the Caribbean, aiid othct Tfiffijitf^'Spedesfi'o^jrtipical awl^^flito^&iFacific 
waters. □ /*(v|/^rA 0emospongia0, Pfi»^cf<cW?fe^<ftr. Hflfew^^. iWRJyfct 'J^^ftfA 

y(?/?/M. Kennedy AlohriN A. Ihopcf. hfatim Biolog)^Lcih0rat0y,Qu£<ti^[Cili<fAft4i^m, 

South Brbhcwe 4101, Australia; J3 Au^uxt. 1999 



Toxic reactions trom handlmu manne sponges 
are well documented fbr speeies i if NfoftlmUlrl^. 
Mimwy Li^sodendoryx^ T^dcmia^ and alsCJ 
fccor&d^hiiTi some speciesnbf Mfm>r/o/w a/id 

//^/Z/VA^^'t/ (sec Wilkinson, 1978: Hooper. Capon 
Si Mudder. lOQl; Hooper. 1<^)'^6: Rifkin, 1906). Of 
iJiese to\ic species, ihe niosi noiorioii:-, is Talnnia 
/^rt/5(Duchassaingc^ Miclielotti, 1864) from the 
Caribbean, earninn it tlie name of '&e iponties" 
(de Laubenfels, 1949). De Laubenfels (1949, 
!954) reported that 7^ i^nis was abundant in 
sliallow-waters throughout the West Indies and 
compared its dermatitis etfects to those of 
poiiSOift'lvy {Rkus faxicodoidwn), producing a 
^sotnewhat painful., itcbiQg, biuning feeling, 
testing for several days* (1 "349: 17% 

Tedania igfiis was described subsequently 
ftotti Hawaii and Patau by de Laubenfels (1950^ 
1 954 )y with some hesitation. Tlieir identification 
of these Pacific speeimens \sa-: niilLicnced by its 
SimilaTity of dcrnnUiiis ie;ieiit)n lo of T /e//M' 
Identification was provisional, and Mcv cnn- 
sidering.jth^ geogfa^ hie isolation, de Laubenfels 
su^ested* fbst they snoutd be recognised as T. 
^/j^iSlibspecies pacifica, 

Witiittje possible exception of a casual obser\'- 
ation hy Bergquist (reported in Soutfacott & 
Cifulier, ]071), such demiatitis reactions have 
not been reported trom any other species of 
Tadwiiu. Bergquist informed Southcott &. Cptd- 
Itiir (197 1 ) that sh^ b^^ ceceiv^ skin irritaliti!n$ 
frdra "handling Tedanmml^di^^^w5'--w&jlmj 
but her observation was not aceompanicd by 
identification or description i>f the oftcndxng species. 

Recent collection of a red sponge from Vanuatu 
produced a skin irritation similar to that 
described for T. igi/is. Subsequent taxonomic 



dithering ftom its congeners in spiculation and 
^ktlmf ^tfii^te, Tbi§ paper describes ihc 
material flsr^ji^&iv^^eeies. det^iing. difibrcpe^ 

^UbltO^iCill waters. 

MAfhRIALS AND METUOD^^^ 

Specimens were collected from the iniertidul 
/•oi tc, I icserved iiiitially in 95% ethanol for four 
dnys, ificn transferred to ethanol for per- 
manent storage. Histological teebiiiques for light 
and scanning electron nncroscopy (SEM) follow 
Hooper (1996). Spicule tnorphometnc analysis 
was conducted miug a light microscope md 
4eAQiec9^1vci^^ u^bf^fiwiic^^t^^ teuipiatc drawn 
from a's^&micmm6fer. Spicule fhisasurcmcnts 
are based on 25 S|iicLiIes ofeaeh spicule category' 
lor each individual., and pertain lo ina\imum 
dimension, denoted as range (and mean) of length 

and width. Spivuk me^sur?iji*?jits ^re in 
microractrcfi. 

AbbteviatioTis: ORSTOM. Instilut Franvais de 
Recherche Scientiiique pour le De\ eloppement 
cn Coop(^ra(ion, Cenlrc de Noumea: QM , Oi.icens- 
jand Mmcum, Brisbane; ^^A, Zoolpgischc 
Museum,IMv0skdtvau Amskidfll^^ 

SYSTEMATIC^' 

PORIFERA Grant 
OEMOSPQNCMAJtv SpUa^ 

DEFINITION>.^Bi^Sling, massive or digiiaie 

sponges; chp?rtios«mflL ^fealc^oo predcHiwnanUy 
plumdfenculdte Of tvth pluttios^, 'composed of 

tracts of smooth or splncd sr\ les, or smooth 
iixcax enclosed withui hgiit or moderate spongni 
fxf?tc^r QT wM\ «io viftihle i\\irc% and spicule; 



446 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 1. Tedania (Tedumu) strongslosiyla sp. nov. (halqtypeQM Ci3 15594). A. Holotype. B. scclion through 
peripheral skeleton. C, sttongylote style and D, terminations. E, tyiole and F microspined base. G, lai'ger 
oi^chaete and H, asymmetrical terminations^ Jj-analler onychaete and J, asymmetrical terminations. 



NEW SPECIES OF TOXIC TEDANIA 



447 



merely cemented together with collagen at their 
nodes; ectosomal spicules are tylotes ortomotes, 
usually with basal spination, lying tangentiaJly, 
paratangenlially or erect on the surface, ahhough 
usually not in bundles; microscleres are 
onychaetes; chelae absent (from Hooper & Wieden- 
mayer, 1994). 

REMARKS. Discussions surrounding the family 
are well summarised in Hooper & Wiedenmayer 
(1994). 

Tedania Gray, 1967 

Tedania Gmy. 1867: 520. 

TracPiytedania Ridley, 1881: 122 (type species Trachy- 

tedania spinata Rid]ey, 1881, by original designation). 
Tedaniopsis Dendy, 1924: 366 (t>'pe species Tedwiiopsis 

turbinafa Dendy, 1924, by original designation). 
Paratedcmia Burton, 1929: 441 (type species Ocecmapia 

tantiila Kirkpalrick, 1907, by original designation). 
Oxyfedania Sara, 1978: 61 (type species Oxytcdania bifaria 

Sara, 1978, by original designation). 

TYPE SPECIES. Reniera digitata Schmidt, 1862, by 
subsequent designation (see Koltun, 1959: 154). 

DEFINITION. Massive; ectosomal skeleton 
composed of tylotes ortomotes with microspined 
bases forming tangential or paratangential 
surface tracts; choanosomal skeleton composed 
of styles with smooth or microspined bases, 
producing reticulate, plumo-reticulate, plumose 
or even dendritic architecture; microscleres are 
onychaetes (from Hooper, 1998). 

REMARKS. The synonymy of Tedania follows 
Burton (1932), with the addition of Oxyiedama 
Sara, 1978 proposed by Desqueyroux- Faundez & 
van Soest (1996) on the basis that the genus was 
unrecognisable, conditionally qualifying it as a 
junior synonym. Desqueyroux-Faundez & van 
Soest (1996) further propose retaining Tedania, 
Tedaniopsis and Trachytedania as subgenera. 

Subgenus Tedania Gray, 1867 

DEFINITION. Tedania possessing smooth, 
relatively small, occasionally strongylote styles 
as structural megascleres, and microspined 
tylotes as ectosomal megascleres (from 
Desqueyroux-Faundez & van Soest, 1996). 

REMARKS. Tedania differs primarily from 
Tedaniopsis and Trachytedania in having t>'lote 
(rather than tomote) ectosomal megascleres. 



Tedania (Tedania) strongylostyla sp. nov. 
(Fig. 1, Table 1) 

ETYMOLOGY. Strongylostyla, for the strongylote-like 
ends of the styles which differentiate this species from its 
congeners. 

MATERIAL. HOLOTYPE: QM G3 15594: inlet leading 
to Yeu Metenia Bay (Picot Bay), Hiu (North Island), Torres 
Islands, Vanuatu, 13°05.340'S,166°33.061'E, miet with 
rocky coralline substrate and moderately turbid water 
(about 20cm visibility), 0.3m depth, 22.vii.1999, coll. JA 
Kennedy 

COMPARATIVE MATERIAL. PARALECTOTYPE: 
ZMA POR.2373 Thalysias ignis Duchassaing & 
Micheiotti, 1864 from St Thomas, Caribbean.. 

H.\BITAT DISTRIBUTION. Marine, less than 
Im depth, on rocky coralline substrate and 
partially buried in surrounding sand, occurring in 
moderately turbid water; Torres Islands, Vanuatu. 

DESCRIPTION. Shape. Thickly encrusting, 
amorphous mats, up to 16cm in greatest 
horizontal width and 2cm thick; loosely adhering 
to rocky coralline substrate and partially buried in 
sand, with surface barely protruding through 
substrate. 

Colour. Bright orange-red externally (Munsell 
lOR 6/12), drab greenish-grey in the peripheral 
Choanosome (2.5GY 6/2), becoming lighter 
brownish-grey in deeper regions (2.5 Y 7/2) when 
alive; ethanol preserved material has drab 
milky-orange exterior, grading toward beige 
deeper in the choanosome. 

Oscides. Small, approximately 1mm diameter 
when alive, scattered indiscriminately over the 
surface, commonly apical on short conulose 
projections up to 4mm high and 8mm diameter, 
but also flush with surface; less obvious in 
preserved state. 

Texture. Soft, spongy, compressible, easily torn. 

Surface characteristics. Opaque, with approx- 
imately two-thirds of surface covered by sandy 
silt and fine algal filaments which extend into 
choanosome; lightly rugose, covered with small 
iiTegular ribs, lightly membranous over irreg- 
ularly scattered, minute, subdermal depressions 
commonly about Inim but up to 2mm wide. 

Ectosome. Difficult to detach from choanosome; 
about 60-100|jLm thick; consisting of a tangential 
to paratangential layer of loose paucispicular — 
multispicular tracts of tylotes in whispy, 
dendritic-plumose arrangement, with abundant 
single tylotes and scattered onychaetes between 



448 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 2. Tcckinia iTedaniu) iguis ( Duchassaing & Micheiotli, 1864). (paralectot\'pe ZMA POR-2373). A, 
Paralectotype. B, section through peripheral skeleton. C, strongylote style and D, terminations. E, tylote and F, 
microspined base. Ci larger onychaete and H asymmetrical terminations. I, smaller onychaete and J» 
asymmetrical terminations. 



NEWsrTcihsoF loxu: iLUAm 



tracts: ectojfomal niembtEme appears veiy gtoti'- 
ular and contains tine detritus fragments. 

Chouiiosome. Skeleton consists priniariK of a 
vaguely ascendingplunin-retibulale arrangernenl 
of paucisni^^ikr vacts ufi{ft|Mitsed roai^^y x>t' 
;fttrongylote ^ityte8 mi fi^cr wrnot^s, wnb 

abundant nicgasclcresandmicrosclcrcs scattered 
IwdividualU' hclvveen tracts ; mi'sohvl is giMiiuljn 
ganlainingb(»tli line and larger detritus fragments 

M^att^tcd throughouti fibres abst^iU; brighl 
orange-red larvae, about SOO^xm diameter, (sood* 
IttOn m deeper chcraiWMOme. 

Me&U&cktes. Strongyfete tliin, smooth, 
Slr^btqr ver^ fienfly cUrVediirat tapering along 
entire length; with ^trong\1otc tcnninations tliat 
?|fe tightly (eleseuped (2 1 (H23_S)-304 x 2,5- 
(3.5 I. Ty lutes, smooth, straight, with oval, 
inien>spined,ipiees(2l3-(22S)-240x2-(3.S)->). 

MiaVM'lefes. Onyehaeles, in two size classes, 
wth abundant spinalion. Both larger ( 1 1 S-( 1 XS)- 
220 X I-( 1 .2)-l .5) and smaller onyehaeles 
(43-(55)-l03x0.H0.7)-l} are iisymmemcal/ 
styloid dtic to micmspiiiUEt^on Icsc^dopc ct^d. 

RI MAIvKS. Tcihviia stvoii<j,ylo\ry!a sp. nov. is 
si*|)L-i llcialK' siuiilai lo tlic ( arililK-aii T r,^m\ 
^Duehassaing A: MicheitUti, 1 Srt4 ) m urowih lonn. 

Spicule dimensions (Table I ) and in producing a 
<fermatiti& reaction upon contact with skin. This- 
similarity in Ihcir spicule dimensions is nol 
surprising, since I.ehncrt <^ \ ati Socst ( 1 996: 69) 
state, Tfihmui (Tethnia) from tropical localities 
«ill over hic v\orld display similar ^ipiculation, so 
bat may not be a goDd species criterion'. 
llTcs|"»cciivc of these aimjlarides^ Z sfipf^lo- 
sv^a sp. nov. differs from T> /^nir lix having 
distinctly differentstyjetertntoations and skeletal 
4lfvbitecniTe« 

Tidankft^iff^Nz^ rodescribed comprehensively 

tiy van Soest ( I ^M4 ). If has an irregular renieroid 
choanosomal skeici;il reticulation, whereas 
Tt'dania styoijoylosiyla sp. nov. has a loose, 
vaguely ascending, plunio-retieulate choanosomal 
skielclai arrangement. Sinnlurly„ T strongyhslyla 
$p.. nov. ba& distinctly .&trongylote styles 
compared with the iittfti(idflK<!' Styles of T. Ignis 
fSF.M examination of* the paraleclotype's 
spiculalion is presented in Fi^. 2 foi eoinpai ison). 
.Apart t>oin the single record ot^strcjngylote mod- 
tficaiions of styles observed in a i^ingle Jauvajeau 
daep-xvsiter specimen tentativety assigned to t, 
(T) et: ignis by Lehnert & van Soest (1996), 
difierences in skeletal arranccmcni, spicule 

morphology and d^jun«1 bK%9»grApbtcdl 



(Jisiribuiions support the. lecogniiion <d' /, 

.v// -'//;^i A'.v/r/j sp. nov. asdistifvri ironi 7' /h.'(ja 

Other species of Tedmna Irom tlvc uopical 
Pacific with two scat cbsscs ol oiiydiaeies incfude 
T. dirhaphin Hcniscbcl, 3912, gahipa^ettsia 
Dcs4ueyroiOft-1^a«SnWs!JS.Vati.SdieSt, 1996 and T 

st}on$:^^la hnhc. 19S6-Thc first two species differ 
significantly tV-.»rn T, stm^i^^lostvlu sp. nt)v. in 
havnig sfyles cl [ypical morphology and mesh- 
Qrpe choai^ostimal skeletal stnicture. TecLutut 
WOngyitt Jinhe. 1^86, described from Chmcsc 
waters (.linhe, 1 986) is siniilar to T ,sftmigylosi(yJa 
sp. nov. in its skeletal arrangement and in 
possessing choanosomal i-.Ii ongy Ics, hue as 
observed for /, ignis these spicules cleiU"ly 
represent malfonned styles and do not constitute 
ihe principal uhoani>fiOmal ^piuukstypef TedfNw 
bra'^ffieftsnrM(:ifb^ etal., 2D00 Ito^ Br^italao 
has t'.vi' st/e classes oftmyehaeies hui diflers; 
Irom 7^ i:}ofif:yf(Ks-iyla sp. nov. m havmg a 
suhisodfciyal chi)ai \^ >somal skeleul arrang^nieittt 
similar In lhat of 7 ignis. 

It ispo.ssible that other species of Tahiniu i i.iiv 
also have two si/e classes of onvLhaeles. L-v cn 
though they were oiiginally recorded as liavij!i{ 
ni\|y one. For example, a Second caiegor\ of 
onychaetc was discovered by van Socst ( 1 9S4 ) in 
L igtiLs^ cind in several Tcdamu (TrachyfedtmM/ 
^t^y DesqwcyrotiX-Fgiiw^ 

IW dermatitis reaction experienced ^ ibc 
primary author through contact with T. sit&iit^h 
losn-lu sp, no\. cDnmienced as a tnild itchinig, 
sensation lasting for about five minutes, 
intensi^ing to severe itching, mi Id swelling and 
n?ddening4)f :tbeakin tosUos for three di^, v^iUs 
subscqucntSKiTi l6fisexpcnencedti«ert>ne«wk^ 

The cs'Ienl of the reaction varied betv. .-en 
collectors, ranguig Ironi only Tiiild iIcHitils lu 
i>ioic severe icaciions as descnlH-.j .ii>i.>vc- 
Expcrifflenial agplicauon ot an alcohol preserved 
spocjmen ^fed 10 produce ^Irritationi^ 

ACK.NO W LI DOn. MENTS 

the anlhurs thaok^OKS'IOM lor the opportm^ 
\ty to participate m ihe norfheni Vannata 

biodjversiTA suncy progratn cimducieJ dtiTing 
July. I9'>>; the captain, crew and scienlisf.s of 
"N/O Alis' for iheir assistance in the field , Dr Roh 
van Soest, Zoologische Museum. IJnivcrsitcil 
van Amsterdam, for loan of type material : an* I i>r 
Eduardo Hajdu, Universidade Federal do Rio dc 
Janeiro and UniversidNje de S»t> Paulo, for 
hHujing to our attcniiannewlftGittbicp. 



450 



MEMOIRS OF TliE QUEENSLAND MUSEUM 



TABLE 1 . Comparison between spicule dimensions of Tedania (T.) strongylostyla, T. (T.) strong\da,T. (T.) ignis, 
T. (T.) brasiliensis, T. dirhaphis and T. (T.) galapagensis. Measurements given in ^m, denoted as range (and 
mean). L=length; W=width. 



Species 


Locality 


Styles 


Tylotes 


Large onychaetes 


Small onychaetes 


T, strongy/ostyia 
sp. nov. 


Northern Vanuatu, 
W. Pacific Ocean 


Strongylote styles 
L. 210-(235)-304; 
W. 2.5-(3.5)-5 


L. 213-(228)-240; 
W. 2-(3.5)-5 


L. il8-(185)-220; 
W. ]-(1.2)-1.5 


L. 43-(55)-103; 
W. 0.5-(0.7)-l 


T. strongy'la 
Jinhe, 1986 


Gulf of Tonkin, 
South China Sea 


Typical styles 
L. 190-310; W. 6-8. 
Strongylote styles 

L. 4-l.--Z.l-4, W. O-O. 


L. 201-218; W. 3-4 


L. 126-182; W. 2-3 


L. 50-62; W. 1 


7". ignis 

yUUL.II. Oi. IVlIt-il., 1 OVM' / 

(Paralectotype; van 
Soest, 1984) 


■ 

Jamaica, C-aribbean 
Sea 


L. 220-240; W. 4-8 


L. 210-225; W. 3 


L. 180 


L. 50 


T ignis 

(Duch. & Mich., 1864) 
(van Soest, 1984) 


Caribbean Sea 


L. 202-(248.8)-281; 
W. 4-(6.31)-9 


L. 180-(2:17.l)-248; 
W. 2.5-(3.38)-4.5 


L. 154-(211.1)-247; 
W.0.5-(1.61)-2.5 


L. 30-(64.0)-95; 
W.0.5 


T. cf. ignis (Duch. & 
Mich.. 1864)(Lehnert 
&van Soest. 1996) 


Jamaica, Caribbean 
Sea 


L. 250-300; W. 9-11 


L. 2 15-240; W. 3-4 


L. 215-240; W. 3-5 


L. 35-70; W. 1 


T. ignis pacifica 
(Duch. & Mich., 1864) 
(de Laubenfels, 1954) 


Hawaii, Central 
Pacific Ocean 


L. 160-210; W. 6-8 


L. 180-210; W. 3-4 


L. up to at least 200; W. 1-2 


/, ignis pacifica 
(Duch. & Mich.. 1864) 
(de Laubenfels, 1954) 


Palau, W. Pacific 
Ocean 


I. .225; W. 3.5 


L. 245-260; W. 5-6 


L. <2I5; W.<1 


T. brasiliensis 
Mothes et al., 2000 


Chilean Coast, E. 
Pacific Ocean 


Slrongyles 
L. 151-228 


L. 151-257 


L. 95-200 


L. 40-78 


T. dirhaphis 
Hentschel, 1912 


Arafura Sea 


L. 218-312 


L. 224-248 


L. 200-312 


L. 40-112 


T. galapagensis 
Desqueyroux-Faundez 
& van Soest, 1996 


Galapagos, E. Pacific 
Ocean 


L. 192-246; W. 6-7 


L. 179-234; W. 3-4 


L. 173-205; W. 2 


L. 61-93; W. 0.5-1 



LITERATURE CITED 

BURTON, M. 1929. Porifera. Part 2. Antarctic sponges. 

British Antarctic ('Terra Nova') Expedition, 1910. 

Natural History Report, Zoology. 6(4): 393-458. 
1932. Sponges. Discover\- Reports 6: 237-392. 
DENDY, A. 1 924. Porifera. Part 1 . Non- Antarctic sponges. 

British Antarctic (Terra Nova') Expedition, 

1910. Natural History Report 6(3): 269-392. 
DUCHASSAING DE FONBRESSIN, P. & 

MICHELOTTI, G. 1864. Spongiaires de la mer 

CaraVbe. Hollandische Maatschappij Der 

Welenschappen, Haarlem. Natuurkundiee 

Verhandelingen(2)21(3): 1-115. 
DESQUEYROUX-FAUNDEZ, R. & SOEST, R.W.M. 

van 1 996. A review of lophonidae, Myxillidae and 

Tedaniidae occurring in the South-East Pacific 

(Porifera: Poecilosclerida). Revue Suisse de 

Zoologie 103(1): 3-79. 
GRAY, J.E. 1 867. Notes on the arrangement of sponges, 

with the description of some new genera. 

Proceedings of the Zoological Society of London 

1867: 492-558. 
HENTSCHEL, E. 1912. Kiesel- und Homschwamme 

der Aru und Kei-Inseln. Abhandlungen 

Senckenbergiana naturforschende Gesellschaft 

1912:295-448. 
HOOPER, J.N.A. 1996. Revision of the Microcionidae 

(Porifera: Poecilosclerida: Demospongiae), with 



description of Australian species. Memoirs of the 
Queensland Museum 40: 1-626. 
1998. Sponguide: Guide to sponge collection and 
identification. (Queensland Museum: Brisbane) 
(http://www.qm.qld.gov.au/nature/explorenature/ 
spong.pdf). 

HOOPER, J.N.A., CAPON, R.J. & HODDER, R.A. 
1991. A new species of toxic marine sponge 
(Porifera: Demospongiae: Poecilosclerida) from 
Northwest Australia. The Beagle, Records of the 
Northern Territoiy Museum of Arts and Sciences 
8(1): 27-36. 

HOOPER, J.N.A. & WIEDENMAYER, F. 1994. 
Porifera. Pp. 1-624. In Wells, A. (ed.) Zoological 
Catalogue of Australia. Vol. 12. (CSIRO Australia: 
Melbourne). 

JINHE, L. 1986. Sponges as marine fouling organisms 
in China waters I. Studia Marina Sinica 26(2): 
76-116. 

KIRKPATRICK, R. 1907. Preliminary report on the 
Monaxonida of the National Antarctic 
Expedition. Annals and Magazine of Natural 
History (7) 20: 271-291. 

KOLTUN, V.M. 1959. The Comacuspongida of the 
northern and far eastern seas of the U.S.S.R. Pp. 
1-236. In Keys to the Fauna of the U.S.S.R. 
Published bv the Zoological Institute Akademia 
Nauk SSSR Vol. 67 Moscow: Izdatel'stvo 
'Nauka' (in Russian). 



NEW SPECIES OF TOXIC TEDANIA 



451 



LAUBENFELS, M.W. de 1949. Sponges of the 
Western Bahamas. American Museum Novitates 
1431: 1-25. 

1950. The sponges of Kaneohe Bay, Oahu. Pacific 

Science 4(1): 3-36. 
1954. The sponges of the west central Pacific. 

Oregon State Monographs, Zoology 7: 1-306. 
LEHNERT, H. & SOEST, R.W.M. van 1996. North 

Jamaican Deep Fore-Reef Sponges. Beaufortia 

46(4): 53-81. 

MOTHES, B., HAJDU, E. & SOEST, R.W.M. van 
2000. Tedania brasiliensis new species (Demo- 
spongiae, Poecilosclerida, Tedaniidae) from 
Brazil, with some remarks about the genus 
Tedania in the tropical southwestern Atlantic. 
Bulletin of Marine Science 66(1): 1-11. 

MUNSELL, 1977. Munsell Color Charts for Plant 
Tissues. 2nd Edition (Munsell Colour: University 
of Wisconsin). 

RIDLEY, S.0. 1 88 1 . XI. Spongida. Homy and siliceous 
sponges of Magellan Straits, S.W. Chili, and 
Atlantic off S.W. Brazil. Proceedings of the 
Zoological Society of London 1881: 107-137, 
140-141. 

RIDLEY, S.O. & DENDY, A. 1886. Preliminary report 
on the Monaxonida collected by the H.M.S. 
'Challenger'. Annals and Magazine of Natural 
History (5) 18: 325-351, 470-493. 

RIFKIN, J.R 1996. Phylum Porifera (sponges). Pp. 
340-344. In Williamson, J.A., Fenner, P.J., 
Burnett, J.W. & Rifkin, J.F. (eds) Venomous and 



Poisonous Marine Animals: Medical and 
Biological Handbook. (University of New South 
Wales Press: Sydney). 
SARA, M. 1978. Demospongie diacque superficiali 
della Terra del Fuoco (Spedizioni AMF 
Mares-GRSTS e SAI). Bolletin dei Musei e degli 
Instituti Biologici dell 'Universita' di Genova 46: 
7-117. 

SCHMIDT, E.O. 1862. Die Spongien des Adriatischen 
Meeres. P. 1-88 (Leipzig: Engelmann). 

SOEST, R.W.M. van 1984. Marine Sponges from 
Cura9ao and other Caribbean Localities. Part III. 
Poecilosclerida. Studies on the Fauna of Curasao 
and other Caribbean Islands (199): 1-167. 

SOUTHCOTT, R.V. & COULTER, J.R. 1971. The 
effects of the southern Australian marine stinging 
sponges, Neofibularia mordens and Lisso- 
dendoryx sp. The Medical Journal of Australia 2: 
895-901. 

THIELE, J. 1905. Die Kiesel und Homschwamme der 
Sammlung Plate. Fauna chilensis. Zoologische 
Jahrbucher, Jena Supplement 6: 407-496. 

TOPSENT, E. 1928. Spongiaires de F Atlantique et de la 
Mediterranee provenant des croisieres du Prince 
Albert ler de Monaco. Resultats des Campagnes 
Scientifiques Accomplies sur son Yacht par 
Albert ler Prince Souverain de Monaco 74: 1 -376. 

WILKINSON, C.R. 1978. Description of two Demo- 
spongiae, one being toxic from the Great Barrier 
Reef Tethys 8(3): 267-270. 



RESOLVING THE 'JASPIS STELLIFERA' COMPLEX 



JOHN A. KENNEDY 

Kennedy, J. A. 2000 06 30: Resolving the 'Jaspis stellifera' complex. Memoirs of the 
Queensland Museum 45(2): 453-476. Brisbane. ISSN 0079-8835. 

Five species included in synonymy with Jaspis stellifera (Carter, 1879) (Coppatiidae) by 
authors were re-evaluated from type material for the first time since originally described. 
Original descriptions were found to be mostly incomplete, with subsequent synonymies 
excessive. Only two {StelIetti)iopsis coriacea Carter, 1886 and Stellettmopsis purpurea 
Carter, 1 886) are retained in synonymy with J. stellifera. Stellettinopsis carteri Ridley, 1 884 
is synonymised with Rhabdaslrella globostellata (Carter, 1883) (Ancorinidae). 
Stellettinopsis hitca Carter, 1886 is retained in Jaspis and S. tuberculata Carter, 1886 is 
referred to Stelletta, both reinstated as valid species. Two new species (Asteropus radio- 
crusta.. Jaspis cristacorrugatiis) were described from one misidentified syntype of S. 
tuberculata Carter, 1886 and other new material superficially resembling ^J. stellifera'' of 
authors. Tropical and subtropical specimens of 'J. stellifera\ comprising much of the 
material described in the marine natural products literature, were found to have been 
misidentified specimens of/?, globostellata^ apparently lacking triaenes. With the exception 
of 7?. globostellata, which has a distribution throughout much of tropical and subtropical 
hido-Pacific, species were found to have restricted distributions in Victorian and Tasmanian 
waters. DPorifera. Demospongiae. Coppatiidae. Ancorinidae, Jaspis stellifera. taxonomy, 
new species, revision. Australia. 

John A. Kennedy, Queensland Museum, South Brisbaiie 4101, Australia {e-mail: 
JohnK@,qm.qld. gov.au) : 20 October, 1999. 



The name 'Jaspis stellifera' (Carter, 1 879) is 
widely cited in the marine natural products 
chemistry and other biological literature (e.g. 
Ravi etal., 1981; Ravi & Wells, 1982; McCaffrey 
& Endean., 1985; Fuerst et al, 1999; Wilkinson 
et al., 1999). This species has an alleged dis- 
tribution throughout the tropical West Pacific 
region, also reportedly occurring along Aus- 
tralia's NE and S coasts, extending into Victoria 
and Tasmania (Hooper & Wiedenmayer, 1994). 
Chemical investigations of various populations 
of 'Jaspis stellifera'' discovered two molecule 
types: cyclic peptides and malabaricane-type 
Iriterpenes (van Soest & Braekman, 1999). 
However, van Soest & Braekman (1999) concur 
with Fusetani & Matsunaga (1993) that cyclic 
peptides are most probably products of various 
microsymbionts, since similar compounds have 
been isolated from many orders of Demospongiae, 
cyanobacteria and ascidians. Conversely, mala- 
baricane-type triterpenes have been reported 
from specimens of V. stellifera' from Fiji and the 
Great Barrier Reef (Ravi et al., 1981; Ravi & 
Wells, 1982), and are suggested as good markers 
for Stelletta s.l. (including closely related 
RhabdastreUa) (van Soest & Braekman, 1999). 
Consequently, van Soest & Braekman (1999) 
proposed that specimens of stellifera'' 



containing malabaricane triterpenes belong to 
Stelletta, lacking triaenes, and not to Jaspis, 

Jaspis stellifera was erected by Carter (1879), 
as Amorphina stellifera, for material from 
Tasmania. Within the following decade, several 
morphologically similar species were described 
from Australia, including Stellettinopsis lutea 
Carter, 1886b, S. tuberculata Carter, 1886a, S. 
coriacea Carter, 1886a and S. purpurea Carter, 
1 886b, from Victoria, and 5. car/m Ridley, 1884, 
from Torres Strait, N Queensland. Shaw (1927), 
under the direct supervision of Maurice Burton of 
the BMNH, synonymised all these species into 
Jaspis stellifera. Subsequently, V. stellifera' y^^s 
described from the Low Isles (Burton, 1934) and 
Heron Island (Bergquist, 1969) on the Great 
Barrier Reef, apparently filling the gap in 
distribution between Victoria in the south and 
Torres Strait in the north. In Bergquist's (1969) 
remarks, however, she disputed Shaw's (1927) 
inclusion of S. coriacea and vS". purpurea in the 
synonymy of Jaspis stellifera^ based on two 
inconsistencies in the published data, both of 
which are demonstrated here to be invalid or 
unsupported (see remarks for J. stellifera, 
belowO- Nevertheless, Bergquist's (1969) revised 
synonymy for J. stellifera was subsequently 
adopted by Wiedenmayer (1989), who examined 
whole type specimens only superficially, and this 



454 MEMOIRS OF THE QUEENSLAND MUSEUM 






1, 2,4,5 ''^^3^ 



FIG. I . Known distributions of species described in this paper. 1 = Jaspis stellifera; 2 = Jaspis lutea; 3 = Jaspis 
cristacorrugatus; 4 ^ Asteropus radiocrusta; 5 = Stelletta tuberculata; 6 = Rhabdastrella globostellata. 



decision was perpetuated in the Porifera volume 
of the Zoological Catalogue of Australia (Hooper 
& Wiedemnayer, 1994). 

In light of the probable widespread misident- 
ifications of tropical West Pacific specimens of 
'J. stellifera\ it is appropriate to re-evaluate this 
species complex, to produce a better infonned 
synonymy and a revised distribution of species 
within this complex. Thus it was necessary to 
redescribe comprehensively all type material, given 
that most original descriptions of nominal species 
were incomplete, illustrated poorly and, as 
discovered here, often incorrect. This paper re- 
evaluates all Jaspis species described previously 
from Australia, since they are all contained in the 
V. stellifera' complex, re-examining key voucher 
material and relevant type specimens, and 
providing a revision of species and their 
corresponding distributions. 

MATERIALS AND METHODS 

Histological techniques for light microscopy 
and scanning electron microscopy (SEM) follow 
Hooper (1996). 

Morphometric analysis of spicules was 
conducted using a light microscope and camera- 
lucida, with reference to a template drawn from a 
stage micrometer. At least 25 spicules of each 
spicule category were measured in all specimens 
(except where noted). Measurements refer to 
length and width of monactinal spicules, rhabd 
length and clad length of tetractinal spicules, and 
diameter of astrose microscleres. Measurements 
refer to maximum dimensions of each spicule. 



denoted as size-range (and mean in parentheses) 
for each spicule type. All measurements are 
given in micrometres unless stated otherwise. 
Centrum percentages for each aster type were 
also noted. 

Conventional morphological terms follow 
Boury-EsnauU & Rutzler (1997). 

Abbreviations. AIMS, Australian Institute of 
Marine Science, Townsville; BMNH, The 
Natural History Museum, London; LMJG, 
Abteilung fiir Zoologie am Landes-museum 
Joanneum (Landes Museum Jubileum Graz), 
Graz; NTM, Northern Territory Museum of Arts 
and Sciences, Darwin; GBR, Great Barrier Reef, 
Queensland; NCI OCDN-, United Sates National 
Cancer Institute, Coral Reef Research Found- 
ation shallow water collection contract, Chunk 
State & Republic of Palau, (1992-present); NCI 
Q66C-, United States National Cancer Institute, 
Austrahan Institute of Science shallow water 
collection contract (1984-91); ORSTOM, 
Institut Fran^ais de Recherche Scientifique pour 
le Developpement en (Cooperation, Centre de 
Noumea; QM, Queensland Museum, Brisbane. 

RESULTS 

Changes to the synonymy of Jaspis stellifera, 
extending from the work of Shaw (1927) to 
Bergquist (1969) and the present study, are 
presented in Table 1. These data propose major 
changes to species groupings within the 'J. 
stellifera' complex. Figure 1 summarises the 
revised known distributions for species included 
in this complex. 



Kh^OLVmG THE JASPlS^STELUFERA COMPL^:X 



45* 



t^BLE 1 . Changing laxononiic status of the ^JaspiiMUifa-a' complex. Bold type = species considered valid; 
toewesan parentheses are considered synonyms of the valid species precedingj • iwiginally syntypeS ibr 
mtHnimpai^ tuhiifculata (Carter); 3 - Identified by Bergquist { 1 ) a.s ktpis 
Oreal Barrier Reef; ^ - included to avoid possible confusion with .A/v/i/s l^iu-u. 

purpureii Carter. 1886b; Stdtemif^i^miea Csictm^ Xji^tbvStettetHnepsis carteri Bi^s^^ 18S4; Stellemu^ja$ 

tuhen uiata Carter, 1886a 

Shaw i 1927) (1 s^cek^y:^tg^ffsiteii^Fa(^tit ^^jr/HS£ori^wJ4rAi^i^^ J^plK 

m herciiiatii) '_ [ [ _ _ 

J asp is purpurea _ 

L unvnt LL'-isignaicOl (7 Species): JiiSpfS ^ftelUfera is', n. Ja^ji^Ls u/riaLto, Jasjnsriufpurca )■ Jaspis iurca; St^etta 
tubervulalul Asteiopua mdiocrusta sp. nm.': Cretin 9p.-; Hhuhtlti&treUa ghoostdtafu ^ (syn. Jttspij^ carteti)iJ^pui 



SYSTEMATICS 

PORlFHRAGram 
DHMOSPONGIAF. SoDas 
T biKACTlMOMORPHA Levi 
ASTIipPJHpRtiM Levi, 1973 

]":l-:FI>jiTION. Encrusting to niassiv^- -rrvvih 
iuiTns; mcgascleres only oxeas turming a 
4Xtt|iUsed, vaguely radial chowosomai skeleton; 
nicgascleres also form a tang^ptial layer in 
eclosome^ triaenes abseiit;trfeix)sd|^s^flSters 
(never slerrasters ), sometimes it^idasters 
(modified from Hooper Wiedenmayier, 1994). 

REMARKS. A surnnuu"} ot" synonymies and 
di:^ussion of the family iire pio\ itied by HunixT 
Sir Wiedenmayer (1994). llajdii & van Soesi 
(1992) suggested thai the absence of triaenes is n 
^pect diagnostic character Ibr the family ai\d 
proposed that the concept of Coppaliidae be 
retained provisioncdiy, pcndmjL; mote detailed 
rc-evaluation of its pr{^bable poly phyiclic nature. 

Jaspis Grayv 1^67 

tSyppittm Sollas. 206 (l)pc species: St^tl^mfntts 

conaa'tf Carter. 1 886. by original designation), 
,4MK/ffcplus Sollas, 1888: 416. 422 iT\nc species: 

,'i\>rnpc;ifu\ pnlrhiT Siil|;is. I XXK. Ii\ iiionoi;. p> i 

"H PE SPEC lB^i, iioa johimmi Schmidi, 1862: 78, by 

DfriNlTTOH. Coppattiaae with wsteis as 

microsc teres. 

KCMAKKS Lendcnfetd (18%) demonstrated 
that Astropepltts pitlcher Sollas. IXSS (type 
species /Uiropcplus) was synonyjiious with 
Koajohnstomi Schmidt, 1862 (type spticie&of 
Jaspis)* bui mititakenly placed it in XcHospOh^ia 
Cray; l858 (tamUy fcthyidae). in recognismg 



Coppadas Solhis, ISSS, reiected Gray's generic 
name Jospts on tlie basis tiiat ii was ot" no 
scieotiJjc vitlue, and dtt:n laicr reinsiaiing./av/;/5 
as a valid genus (Topscnt, 1904 ). In ratifying this 
later decision, Dendy (1916) again synoiiyitiU^d 
Coppatias' \N\{\\ Jaspis, the scmor name. 

Hajdu Si van Soesi ( 1 9V2) quesiioned whetlicr 
at not Jaspis coastituted a monophylcUc 
assemblage, since two species groups were 
recogn isablc based on the presence or absence of 
'microxeijsV However, this is not accepted Here 
as these 'niicroxeas'are considered to be a small- 
ci category of oxeole megaselercs. The e.xi.Htencc 
of .eraded oxcote si/c-distribuii(-ns with iniet- 
meSiate si^e categories alien presents diCtlcuiiifs 
in dif^rentiatizi^ between 'Sm<dler('mkfO]^2k^) 
and larger oxeotes. Henee, the pre.scnt concept df 
Jaspis retains bolli assemblages, althougli i1 is 
LiektiuvvIedgL-d (bal lurlhei work is required to 
reconcile the taxonuniic sigiulipance of smaUec 

Jaspis steOtflBfiHi (CSift«f. Wf^i 
(Figs 1, 2, Tabled 

Imorfih/nu .stcHiJt'i'o CcAHiT, 1879: ^44 , 

S;i!;cf!;i!op.si\ .^Uilt/cra; Ridlo . 18X4. 47T» 

Ci>r(>iiiius M€tlifcni\ Solla:=, 18X8; m 

JaspiK sU-Hiffiv. Shau 1 *)?'^- 422. 

not Jusph; sIc'IlifcTu: I5urion. 0*34: 522- 

Siel/enirutp\/s aniitcca L jrtcr, IsSOar 126.- 

CoppiUias L'ttrkici'us: Soll.-jy. iKSS: 207 

Jaspis ctwccj; I loopcr & Wiedaimara, 1^94; 143. 

StelienirtupMf, putptuva Caner, 1 8tt6b; 4d9. 

M^piijrUrpuriux'yWS^^ 143. 

MATERIAL. HOLOnTE: HMNHI869.L22JJ5 (diy); 
lasmanta (alktci ma|r}cBd'1&rIA-C8TtdC AO 
i lolutvpc of St^kttmcp^s chrfmm '^^it&kl tSMi 
BMN|'msS6.12.15.441 rdr>): IVm Phillip Heads, ViictwiBi 
Holorv po of StelleWnopsis purpurea Carter, I86fib 
BMNlll886.i2.is.5i (\vli). WestempoTt Bay, Vicioria 
OTHBR M'vITRlAl : BMNII unregistered (second 
spficimen m^neeoniaineraff halQ[^and.aIs^ 



456 



MEMOIRS OF TllE QUEENSLAND MUSEUM 




FIG 2. Jaspis stellifera (Carter, 1 879) (holoivpe BMNH1869 J-2Z25). A, holotypci B, holqlype^jf^ e^Pr-itt^ 
BMNH 1 886. 1 2.1 5.44 1 ; C. holotype of 5. pitrpurea BMNHl 886.12.15^1 (arroWTn4itmiS*Sc&Wf>ft o*'^^^ 
D, section throt^ peripheral skeleton; E, oxeas; F, oxyaster. 



RESOLVING THE JASPIS STELLIFEM COMPLEX 



457 



TABLE 2. Comparison between present and published 
descriptions of Jaspis stellifera. Measurements in 
^m, denoted as range (and mean) (N=25). L = length; 
W = width; D = diameter. 





Oxeas 


Oxy asters 


Holotype 

BMNH 1 869.1.22.25 


L 113-(388)-726 
W3-(12)-16 


D ll-(15)-22 


Holotype (Original de- 
scription Carter. 1 879) 


L 725; W 17 


D 17 


BMNH unregistered 
Bowerbank collection 
Carter no. 315,E.h,19 


L 120-(354)-730 
W3-(10)-16 


D7-(12)-]6 


BMNHI886.12.15.441 
holotype of J. coriacea 


L62-(265)-89S 
W2-(8)-21 


D7-(ll)-13 


BMNHI886.12. 15.51 
holotype of./, purpurea 


L53-t347)-781 
W3-(10)-18 


D7-(10)-12 


NMV F5193 
(Wiedenmayer, 1989) 


L 89-(364.8)-682.8 
W2-(9)-]8 


D8-(12)-15 



with Carter no. 315E./7.19) (drv): South Australia, 1859, 
J.S. Bowerbank collection. COMPARATIVE 
MATERIAL: Holotype of Vioa johmtonii Schmidt 1862 
LMJG 15648/0 Sebenico, Adriatic Sea. S>Titype of V. 
johmtonii van Schmidt, 1868 LMJG 15256/0: Sebenico, 
Adriatic Sea. 

HABITAT DISTRIBUTION. Subtidal to 37m 
depth, amidst dense algal growth; Westemport 
Bay and Port Phillip Heads, Victoria; Erith 
Island, Bass Strait; Tasmania. 

DIAGNOSIS. Irregularly lobate-massive; dull 
pinkish purple-brown alive; surface optically 
smooth, even, unomamented; ectosomal skel- 
eton consisting of a densely packed tangential 
arrangement of oxeas largely obscuring micro- 
scleres; choanosomal skeleton of singular and 
loose bundles of oxeas in confused arrangement, 
with scattered oxyasters; oxeas in wide size- 
range (length 53-(339)-898, width 2-(ll)-21), 
microspined oxyasters (diameter 7-(12)-22). 

DESCRIPTION. Shape. Holotype massive, 
amorphous, irregularly lobate, may envelop 
other materials such as algae, shells or detritus. 
Height 4.7cm, width 2.6 X 2.5cm. Other 
specimens up to 5.5 x 4.5 x 3.5cm. 

Colour. Fresh specimens described as dull-purple 
to brownish-drab (5RP 5/6-8) (Wiedenmayer, 
1989); dry holotype creamy-white (Munsell 2.5Y 
8/3) with dark-pink (2.5R 5/4) difiuse patches 
scattered over surface, with greyish-beige (7.5YR 
8/2) choanosome; ethanol preserved specimen 
dark greyish-purple (5RP 3/2) throughout. 

Oscides. Not visible in dry holotype but Carter 
(1879) describes 'vents in pit-like depressions'; 
one incomplete specimen (BMNH1886.12.15.5 1) 
has a single apical depression approximately 4mm 



deep and 11mm in greatest width, containing 
several oscules up to 3mm diameter. 

Texture. Firm, compressible, leathery. 

Surface characteristics. Opaque, optically smooth, 
even, imomamented. 

Ectosome. Thin, approximately 400-700 thick; 
skeleton composed of oxeas in confused arrange- 
ment, with oxyasters scattered throughout but 
largely obscured, and with clumped pinkish- 
purple pigment cells scattered over surface. 

Choanosome. Permeated by canals approx- 
imately 0.15- 1.10mm diameter, with smallest 
canals near periphery; skeleton comprised of 
loose, multispicular bundles of oxeas in confused 
arrangement, with abundant scattered oxeas and 
oxyasters; mesohyl contains clumps of scattered, 
dark-pinkish pigment bodies approximately 2-8 
diameter, and spherical translucent and tt*ans- 
parent bodies. 

Megascleres. (Refer to Table 2 for spicule dimen- 
sions) Oxeas in one wide size-range, t}'pically 
curved over their entire length; variations rare but 
include straight, lightly flexuous, singly and 
doubly bent, and styloid forms. 

Microscleres. (Refer to Table 2 for spicule 
dimensions) Oxyasters with approximately 
10-20 fme, lightly tapering rays with recur\'ed 
microspines on distal two-thirds of rays, centrum 
approximately 13% of spicule diameter; 
variations rare, but include vestigial spination 
over flill ray length, to exaggerated clumping of 
microspines on distal portion of ray, thus vaguely 
resembling tylote terminations. 

REMARKS. The BMNH specimen box contain- 
ing the holotype includes two specimens. Only 
one specimen is pierced by a tag bearing the 
registration number BMNH69.1.22.25, along 
with the locality 'Van Diemen's Land' (Tas- 
mania). The other specimen is from the J.S. 
Bowerbank collection and comes from 'Southern 
Australia'. Carter gave both specimens his 
number '3 15, E, /?, 19'. While both specimens are 
confinned here to belong to J. stellifera, only the 
specimen bearing the BMNH tag is taken here to 
be the holotype. 

Little information is added here to the original 
description except for some detail regarding 
spiculation. As previously noted by Wieden- 
mayer (1989), the oxeas have a wide size-range, 
with little concordance between length and width. 
While smaller oxeas are relatively abundant, the 
presence of many intermediate sizes precludes 
recognising more than one variable size-class of 



458 MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 3. Jospin tutea (Carter, 1886) (lecto^ype BMNH 1 886,12. 1 5.356). A, kcto^pe; B, paraJectotype 
BMNH1886.l2J5,d3 (arrows indicate pos&oti of specimen on Substrate); C section ti^tig^ petipheml 
skeleton; D, large oxea; E> small oxea; F, oxyas^. 



oxeas. SEM examination revealed that miero- 
spination of the oxyasler rays is commonly recLined 
toward the centrum. Spicule measurements 
undertaken hejre confirmed Wifsdenmayer's 
(1989) suspicion that Cstrter provided only the 
measurements for the largest spicules, ignoring 
the range of spicule sizes actually present. 

Stellettinopsis coriacea Carter, l8S6a and S. 
purpurea Carter, 1886b are retained here in 



synonymy with./, stellifera (C^'Sty 1879), based 
on re-examination of all type specii|ieii9, 
revealing that they are morphologically identical 
in virtually all respects. This partially ratifies 
Shaw's (1927) synonymy for X stellijera, and 
contrasts to Bergquist's(1969)revisedsynonymy. 
While Shaw's synonymy was excessive, 
Bergquist's difficulty in accepting the inclusion 
of & coriacea zSi^S^purpureayfd&hdSQd on two 



KLSOLVlNCi rilt JASPIS STELUf^ERA COMPLfc^ 



H59 



sopposed ineMBistendes in the -pdblished data, 

hnlh of whidi are invalid. Firstly, R'..T.uquit;t noted 
ihut Shaw's specimen reportedly did not have 
alters: secondly, she slated that 'Jaspis' ccriacca 
and ,/. ptopurea have iwo distinct calegorieji^Qr 
asters, a feature nevci* noted in ,/ sft2ll^era\ t!0 
this debate, it is iinimporlani whether or not 
Shaw's specimen had asters because it was not 
the ty-pc. Caller's ( 1879) original description of 
(hehoUHypo clearly described only one category 
of aster, confirmed here from rc-cxamination of 
typCTnaieml. BeiS^ui;it'&( 1 969) second point of 
aifipuCifttiDn* h ^sb unsupported since original 
descripiions of.S' coy'iacx^a antl S parpm-i'a niso 
include only one category of aster, rather tlioji 
two, also pQpfirni«<JJ«)nVTC-exaniit»ation ofiype 
material, 

Original desciiptions of S. ajt uicea and S. 
pnrpitreu TVport the presence of 'raicroxeas', 
'\vhcix:a9riD-cXBminatio^i)f|ypet^ateri^ reyeal- 
ed YhAl, ^ d«Mdb6d ^mty Iht^ ttiicro^eas 

Tppresent onTy^ smaller examples of a wide si/c- 
Tange of oxcas, with many intermediate sizes. 

ComparisonbtfhVeen J. .sfellifem and the type 
Species of Jaspis {i'ioa /ohnsfitnii Schmidt, 
1862), includitig both type material and pub- 
lished descriptions (Dcndy, 1916; Biu"ion& Rao, 
t932) confirms thfl^</. sielHferu is a true Jasspis. 
Both J. johnsfanik niid X stellifera have u 
t^trtgcntial cciosomc emnposcd ofoxcotc spienle:^, 
and a choanosome coniaining oxeoles in con- 
fused and agLicly radial arrangement. l:!uasierN 
are oxyasters, some with rays thai are mmulely 
micaxJspincdCafeatture not described previously 
fbr the type specif). Ja$pi:i.JohfJsionU difiE^s in 
having a bimods^l 3»!e'dfstrifnifion of nit^OvKb 
ihe larger si/c-class primarily restncTed to lhi6 
i;hoanos(iirie'), v\iiereas J. sft'lltfcya haj^ a wide, 
uainioilal size-dislribution. Irrespective, both 
clcat jy belpt\g to Pjesei^j cpncepi of jE^^is, 

-flffeiP^SJiribullons remaining unresolved. 

Examination oi' % alide oi Burton's (1934) 
specimen (BMNHT93&.RI5.S6) identified as 

\kispis s/ellifcra' Irom tlie Low Isles,. GBR, 
I'iivealed that it too had been misidcntificd 
(Hooper et af. I'^VS); present sludy). Burlt»n's 
specimen clearly difliiars Irom J. stdlifera in 
laddrig h distinof i^li3KS6me and in having two 
categories of aslcrs, one being slightly bipolar 
and resembling dit>laslers or short spirasicrs. Cnoss 
morphological dilTcrenccs cannot be commented 
on here since only a slide was available for cx- 
janmation^ and Burton diil.i>i0A tmbtfeh Amy 



descriptive detail regarding the specimen. It may 

ucll -epreseni a new species, smcc it docs not 
coircs|>i(r*d io any Jn\pf.\ species detiCribtd 
pre'» nuiriy irom Australian waters.. 

Wiedenmayer's (198*^) description ol' a 
specimen of./ stdUfha from southern Auslralia 
■mcluded only superlicial comparison ul'his mat- 
erial with various lype specitnens (y\mnrphi*U} 
stclfifiro, SU'/l'/ffinojy^is n/hercH/ufn and S. 
lufed). Because of circufuatancea preventing htm 
from examining sltdeS'^f theSe types, he wws 
unable to reveal the di^tmctive nature of each of 
these species. This is discussed furtlier below, 

jMspist Intea (Carter, 1886) 
(Figs 1,3, Table 3) 

MAI t:R.I Al I J .( ■ rOTYPH: P.MNI i 1 886.12. 1 5.356 (wel): 
Wostcnipdii X'ictoriii coll LB Wilson (*AR A- 

i.l-r |( J P,-|r. li\.]Nl-llKS6.12.l.\'H (Uiv): VvVMCitip. >fl 

liay. \ ictona, cvU. J.B. WUsorj. OTlM NiAIERL^L; 
15MNIlW54Xlia56 fslMtrtf -Qifttiiepawd 1^ A. 

Dcndvi. 

KABlfAl PlSiiUBUTlON. Siibudai lo 1 3m 
depiH; Wisisterapent Bay, Vtetorisu 

DlAGNO^>lS. Lobatc-massivc, agglomerating 
'»Mb9tCiM^ OnSf^^f^ Surl'ace lobule> lubercuttfiej* 

ecto^ome drainct, comprising a fine layer or 

small oxeas overlaying taniicnlialK -ananged 
larger oxeas; choanosomal skeleton primarily a 
confuted arrangenictn oi'ovcas, oxeas iti tsvot-izc 
classes (larger oxeas length 1^^0-(516)-712, \^idtb 
6-(12)-l'^; smaller oxeas length -^2-(Sl)-llO, 
widvh l-(3>3), piivruspiped oxyAslei^ (digmeler 

DESCRTPTTON. Shape. Irregularly 1o(>arCi 

agglomeralinu calcareous .subslr4ile>. sand 1IM)I> 
shell fragments. Types incomplete, with largest 
rponlon Scm high. 1 2 x sem wideu' 
Ct>hmn Live a»louration unknown; wel Icetotype 
has tan-brown surikce (Munscll 2.5 Y 5-7/6) wilh 
g()ldL-ii-biov\n L-li()aiu»s*mic ^_.-.5^' ^/R), dry 
paralectoiype has dull-creamy ycUo^V fiurfiicc 
(2.5Y 6-7/4) with dull goldett-y^llow 
choanosome (2.5Y 7, SV 

O.sat/es Nuuicfous, apptoxnnately (),5-8-2jTun 
dimneier. Hush \v^tb, ^ regularly difitnbuted 
over surface. 

Tixrure. Firm, slightly irinhlclctttlieiy.. 
Surface charm, ('.-risdi-.s. Opaque, inerahrRUOMS* 
optically smooth, uneven, witli clusters of lobale 
tuBetel^^ lf^£$;Ulstr^ dtsrrilnKed fiver ^ur^ 



460 



MEMOIRS i)K THK QUEHNSLAND MUSRUM 




notmcludedmCancr's (l8S6b) arigbialidescriptm, butincdudBd!^ 





























1 Wt4; W 15 




n 18 j 



Ectosome, Di&tmci Ihnri choanoscmie. approx* 
imately 400-600 thick; skeleton highly spiculose, 
deribieiy packed, coraprLsed of two layers: ouier 
layer iargfty indistinct, very fine, approMiiiaiely 
?0 thick, with small oxeuii jji conftc^^cd arrange- 
mcnt; iruicr layer comprised of a tangential 
arrangement of both large and small oxA&ft;. 
nqiuifiTaus canals, approximately 200 diameter, 
regularly traverse edosotne; oxyasicrs present 
t»itlaigicl>* obscured by tnegasc^cres. 

CktHtwom Perai«8it«d by lA^ oitwk up to 
llfmh diameter; skeleton composed of a con- 
fused arrangemenl of single small and large 
ox^as. as well as large oxeas in loose pauci- 
spicular (0 nuiliispicular bimdles; oxyasters 
scattered throughout thie i^boasoosoii^ skeletott 
l)ut ftlightly momaVoiiUfont fa CAnAMinfii^. 
Megasclcns. (Refer lo Table 3 for ipiailc 
dimensions) Oxeas, in distinctly bimodal si/^- 
disJribuUon. Larger oxeas slightly CLirvcd over 

entire Jengtii, with v^ry fouitly^tclcscc^d poinb 
imd cn»iasiontdly ^^th s"^^ tclminal niicrospmcs; 
variations rare but ineltidc styloid nu>ditlcaiion!i- 
SittAlUfr oxeas, ungulau:. with 1-2 bends *ind 
lia^aE&teniiinali|9iDs. 

MicmscFeres, (Refer io Table T for spicule di- 
mensions) Ox>'aslcrs, with (t-12 liglnly tapering 
rays that havcci>nical torcc irx ed nueruspmcs on 
distal Iwo-tbinis; pentrum approx^imately 15% of 
.^Pli^ule diameter, 

K^ilARKS. JuspLs lufea is reinstated as a valid 
^p^Ci^ of JaspiSt 4isti|iet &<m J» stellifera in 
several intpcxrtsmt refjpects. Thetcr are hriport^'^nt 

difterences in spicukaion, with t/.-^-fe/e^/ having 
two si/e clashes ot oxcas in a diatincfly bimodai 
si7.e-disiribiition (cf J steUifem fajuving^ aiUli- 
modul. wide ^i/e-raiige or pxeotc spicules). 
Skeletal ditTerences imvSitJ. Ute6 naving a 
more localised disti'ibution of sniallei o\ca>i, 
prnnarily in a distinct ectosume (cf no such 



less priomineDi ectosoiiae). There 
are also differences in colour ^ 
external morphology, with ,/ lute'a 
being golden yellow-brown and 
having a lobate-tubeieuiaie growdi 
form (cf. dull pinkish-purple- 
brown with an unornamented, 
irregularly lobate-massive shape). 

hnportant information provided 
here, ridding to Carter's (1886b) 
original dcscriplion, includes the 
description ot a second size-class 
Oxea (perhaps previously dis- 
missed as ttfW''Cly ^mailer cjc^mples of a 
presumed unimi»da1l l5rze-tfetH6iition of wedw 
i-pieuL':-}. the prr>vision of spicule dimeM>ior-s. 
anil details c>l microspinaiion on oNya^ter rays. 

Jaspi9 crlstucpri-i^Qtus sp. nov. 

H IYMOt.OCiV. i,ii\.\n crisui, ridge; Latin, aom^antf, 
wrinkled; tbi tlic coinigaied osculaj ndgc. 

MATFRIM. HOLOTYPE; QMG3 12071 iNCI 

f^VViC^M') N): VV >.idc orchanncl, in middle of Breaksea 
island. Port Davcy, fa-sniania. ;\usTjaiia, 4_^"I*y.70'S, 
145"57.0()'b, exposed rock slope, houldcrs, walls, gullies, 
kelp, JOm depth, I7.ii.l4gi, coll. AIMVNCI. 
PARATYPn- UMG3 1 2073 (NCI Q6(^C-5 1 5 1 -P): S end of 
Breaksea Island, Port DjVLjy. Tasmani^i. Ausrrnliii, 
43^20.20'S, J 47 57.X0' rocky slope to flat rocky bottom 
wUlma^crevicesaiid^^^ 15m depth, 17aU991jCo]L 

KARrWDlSTRIBUtlON lO I5m depth,iBH 
roclc^' substrate witb IfMl^ gullies and o\itcro^$; 

DIAGNOSIS, Ma»rvfl| 5y6«^e*icaJ to siighiiy 
lobate; leathery, fUin; sbtS-gn^ aUvCk darie-; 
bro^ to golden out of wken tah-fei^owti irt 

elbanot; slightly rugose surface, many small 
(>sti!le::: in furrows between corrugatiuns on v\ide 
o^cLilnr ; idge; dixjinel ectosonie. primarily of 
densely packed smaller oxeas; choanosom^ 
riddle Mr]tbxiin4b| odd skeleteir of single and 
paucispicular bundles of large oxeas in conftised 
to loosely plumose reticulation vaguely ascend- 
ing toward surface, with scattered smaller oveas- 
and oxya^iters between tracts; oxeas ni two sue 
classes (larger oxeas length 290-(45 3 )-629, width 
K9>15J; smaller oxeas length 80-(133)-267, 
-wwth 3-(5)-12}; microspined oxyasters (diam- 

T)V^CiJfTVMu Shape Preserved specimen 
incomplete but when living was nussive, dufe-" 
spberteal) slightly Ipbat*, -with n ibic^y 



RES0LV1>JG THE JASPiSSTELLtFERA COMPLEX 



461 




FIG. 4. Jaspis cri.siacorrugafKs sp. nov. (holoupe 12071 ). A. oscular ridge surface; B» perpfindiCular 

Seption; C, section through peripheral skeleton; D. large oxea; E, small oxea: F, oxy aster. 



corrugated, oscular ridge along the apex (from ectosoine with tan-brown (7.5^11 5/6) membrane 

jdjolographic record, Fig. 8A-B> Height 15cra, surrounding oscules, with ian-bro\\TH7. 5 VR 5/6) 

vdi&i 20>&20(^ wt&n alive and Qouiiplet^ <ihoano$6me in edianol 

Co/r?w;: Slate-grey ectosome (Munscll 2.5V fvl ) Oscules. Many, small osl tiles, approximately 

with lempn-yellqw (.2.5y 8/5) iii jnembrane Q.5iT[im4iametei,pluster^dinlinfiaraTta^ 

sumjunding osicules T^iferft aliVes 1*4 oscides ivijd&:i|i mto-WB h^tm corrugate 

(7.5Y 7/8) to dark-brown (SYR 2.5/2) ectosome, ioiis of oscttlactislge. 
with golden-yellow choanoson^e (7,5 YR 7/8) 

wjien freshy ^ohoi^oim-hrnvm <7*5Yii 4/2) r^octera Veiyfinn,mtbety, wilhleatfiei^ 



462 



MEMOIRS OF THE QUEENSLAND MUSEUM 



Surface characteristics. Opaque, optically smooth, 
uneven, with regularly distributed low rounded 
surface swellings tending toward slightly rugose. 

Ectosome. Distinct from choanosome, 400-1400 
thick, regularly traversed by aquiferous canals 
approximately 200 diameter; skeleton highly 
spiculose, with a densely packed layer of smaller 
oxeas in confused arrangement that largely 
obscures a scattering of oxyasters. 
Choanosome. Riddled with aquiferous system 
canals up to 1 1mm diameter; skeleton of single 
oxeas and loose paucispicular bundles of oxeas in 
confiised to vaguely plumo-reticulate arrange- 
ment; oxyasters present and slightly more abundant 
in aquiferous system linings. 

Megascleres. (Refer to Table 4 for spicule 
dimensions) Oxeas, in distinctly bimodal size- 
distribution. Larger oxeas lightly curved over 
entire length, occasionally fusiform, with acerate 
to slightly telescoped points. 

Smaller oxeas centrally cur\'ed, with hastate to 
slightly telescoped points. 

Microscleres. (Refer to Table 4 for spicule 
dimensions) Oxyasters, with 9- 1 5 lightly tapering 
rays that have recurved microspines primarily on 
distal two-thirds; centrum approximately 
10-15% of spicule diameter. Variations rare but 
include vestigial spination, to clumping of spines 
near terminations, thus resembling t>iotes when 
viewed under light microscopy. 

REMARKS. Even though this material is 
described as a new species, and has not been 
previously synonymised with ./. stellifera, it is 
included here because it may be easily confiised 
with the newly reinstated J. lutea (if the subtle 
differences described here were not elucidated). 
Although J. cristacorrugatus is similar to J. lutea 
in spiculation and choanosomal skeletal 
stnicture, the two are clearly differentiated by 
several important characteristics. The external 
morphology is the most obvious difference 
between the two, with J. cristacorrugatus being 
massive in growth form (cf. J. lutea being 
irregular-lobate), and does not agglomerate 
foreign materials. Further, oscules are grouped 
into furrows across a thick oscular ridge (cf 
regularly distributed over the surface). Significant 
differences in texture also differentiate the two 
species, with J. cristacorrugatus being very firm 
and rubbery (cf crumbly and friable). The main 
differences in skeletal structure is that J. crista- 
corrugatus has an ectosome composed of smaller 
oxeas, lacking the underlying tangential layer of 
oxeas found in J, lutea. 



ANCORINIDAE Schmidt, 1870 

DEFINITION. Growth forms encrusting or 
massive, or more specialised with spherical body 
and long inhalant and exhalant tubes at opposite 
ends (the latter with a stellate, spicular, funnel- 
shaped end); megascleres long-shafted Iriaenes 
(with shaft directed inwards and clads on the 
surface) and oxeas; microscleres euasters and 
microrhabds; without stcrrasters or amphiasters; 
triaenes may be absent or reduced (modified 
from Hooper & Wiedenmayer, 1994). 

REMARKS. Ancorinidae Schmidt, 1870 is best 
kno\Mi under its junior synonym Stellettidae 
Carter, 1875 (Hooper & Wiedenmayer, 1994). 
Hajdu & van Soest ( 1 992) provide an informative 
discussion on the relationship between Ancor- 
inidae and Coppatiidae. 

Asteropus Sollas, 1888 

TY?E SPECIES. Stellettinopsis simplex Carter, 1 879: 349, 
b\' original designation. 

DEFIIMITION. Ancorinidae with oxeas, oxyasters 
and sanidasters to which trichodragmata may be 
added. 

REMARKS. Bergquist (1965, 1968) and Hajdu 
& van Soest (1992) proposed that two species 
groups exist within Asteropus based on micro- 
sclere type. They argued that species with true 
sanidasters may be placed in the simplex' -Wk^ 
complex, whereas those with spiny microrhabds 
should be placed in the ^ sarasinorum' -\\kQ group, 
the latter ascribing the name Melophlus Thiele, 
1 899 and possibly valid at the subgeneric level. 

Asteropus radiocrusta sp. nov. 
(Figs 1,5, Table 5) 

Stellettinopsis tuberculata (in part) Carter, 1 886a; 1 26. 

ETYMOLOGY. Radius, Latin, ray; crusta, Latin, hard 
outer surface of a body; for the radial arrangement of 
megascleres iji the cortical ectosome. 

MATERIAL. HOLOTYPE: BMNHI886.12.15.146: Port 
Phillip Heads, Victoria, coll. J.B. Wilson (originally one of 
three s>"nt\'pes of Stellettinopsis titherculata Carter, 1 886a). 

HABITAT DISTRIBUTION. 3-6m depth; on 
granite boulders with algae; Port Phillip Heads, 
Victoria. 

DIAGNOSIS. Massive, subspherical; ectosome 
forming a highly distinct cortex 2.4-3 .1mm thick, 
of densely packed oxeas in radial arrangement, 
with sanidasters and few oxyasters scattered 
throughout; choanosome with oxeas in confi^ised 



RESOLVING THE JASPIS STELUFERA COMPLEX 



463 



arrangement, with oxyasters and few sanidasters 
scattered throughout; oxeas (length 530-(1063)- 
1730, width 5-(24)-44), inicrospincd oxyasters 
(diameter 18-(25)-33). microspined sanidasters 
(length 9-(13)-18). 

DESCRIPTION. Shape. Massive, subspherical, 
with slightly irregular surface. (Attached to the 
holotype is a second species of sponge that is sub- 
spherical and has a highly conulose, membranous 
surface). 

Colour. Live colouration unknown; beige-grey 
(Munsell 7.5YR 6/2) cortical ectosome with light 
beige choanosome (7.5YR 7/4) in ethanol. 

Oscules. None visible. 

Texture. Harsh, firm, barely compressible. 

Surface characteristics. Opaque, optically smooth, 
even, microscopically hispid and densely 
spiculose. 

Ectosome. Approximately 2.4-3. 1mm thick, 
forming a cortex that is highly distinct from 
choanosome; skeleton comprised primarily of 
oxeas arranged radially in loose muhispicular 
bundles forming an almost continuous palisade, 
with oxea terminations commonly penetrating 
the surface; sanidasters are scattered throughout 
the ectosomal skeleton but are slightly more 
common at the surface, while oxyasters are rare; 
containing abundant subspherical pigment 
bodies approximately 25 diameter. 

Choanosome. Skeleton consists of a confused 
arrangement of both single and very loose pauci- 
spicular bundles of oxeas, with an abundance of 
interstitial oxyasters, but few sanidasters. 

Megascleres. (Refer to Table 5 for spicule di- 
mensions) Oxeas in a single, wide size-range, 
typically cur\'ed over entire length, with lightly 
telescoped terminations; variations rare but 
include styloid fonns. 

Microscleres. (Refer to Table 5 for spicule 
dimensions) Oxyasters, with 7-14 tapering rays 
that have abundant, recurved microspines 
occurring along the entire ray length; centrum 
approximately 12% of spicule diameter. 

Sanidasters, with conical microspines and 
approximately 10-16 rays in 2-4 whirls. 

REMARKS. The holotype of this species was 
originally part of the synt>^pe series of Stellet- 
tinopsis tuberculata Carter, 1 886a. However, it is 
clearly different from the other syntype (now 
lectotype, BMNH1886.12.L5.434) of^". tubercu- 
lata), with the most significant differences 
involving spiculation, skeletal structure and 



TABLH 5. Spicule dimensions of Asteropus radio- 
crusla. Measurements in jim, denoted as range (and 
mean) (N=25). L = length; W ^ width; D = diameter. 





Oxeas 


Oxyasters 


Sanidasters 


Holotype 
BMNHI886. 
1 12.15.146 


L 530-(1063)-1730 
W 5-(24)-44 


D I8-(25)-33 


L9-(13)-I8 



external morphology. Asteropus radiocrusta has 
a single size category of oxeas, as well as 
oxyaster and sanidaster microscleres (cf. S. 
tuberculata which has two sizes of slightly 
flexuous oxeas and triaenes as megascleres and 
only oxyasters as microscleres. These substantial 
differences in spiculation are alone sufficient to 
clearly separate these two taxa at the generic 
level. However, they also differ significantly in 
the skeletal structure of the ectosome, with A. 
radiocrusta having a highly distinct, thick 
cortical ectosome of oxeas in erect to plumose 
bundles, forming an almost completely radial 
pahsade (cf an arenaceous cortical ectosome 
with sand-grains largely obscuring scattered 
oxyasters). In addition, their respective growth 
fonns are greatly different, with A. radiocrusta 
being small, subspherical and lacking surface 
ornamentation (cf massive-Iobale and covered 
with prominent, irregular tubercles and ridges). 

The presence of sanidasters places the present 
species within the '5'//7;/?/t'.v'-like group, as 
described above (see Remarks for the genus). 
Asteropus simplex (Carter, 1 879) is the only other 
species of Asteropus described so far from Aus- 
tralia. Asteropus radiocrusta sp. nov. is similar to 
A. simplex, based on its original description, 
apparently falling within the published geo- 
graphical distribution of simplex (Hooper & 
Wiedenmayer, 1 994), which includes most of S 
Australia as well as New Zealand and the Indo- 
Malay region of the Indian Ocean. However, this 
purportedly extensive distribution of A. simplex 
is dubious since re-examination of a type slide of 
A. simplex from Carter's collection by Hajdu & 
van Soest (1992) revealed that the original 
description was incomplete, failing to recognise a 
second size category of oxyaster as well as the 
presence of trichodragmata. Consequently, A. 
simplex may also prove to contain a sibling 
species-complex, with its junior synonym, A. 
haekeli Dendy, 1905 (taxonomic decision by 
Dendy, 1924), certainly warranting its 
re-evaluation. Nonetheless, A. radiocrusta is 
clearly distinct from A. simplex by the absence of 
both the second size class of oxyaster and 
trichodragmata. 



464 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG. 5. Asteropiis radiocrusta sp. nov. (holotype BMNH1886.12.15.146). A. holotype; B, section through 
peripheral skeleton; C, oxea; D, oxyaster; E, oxyaster ray; F, sanidaster with two whirls of rays; G, sanidaster 
with four whirls of rays. 



RESOLVING THE JASPIS STELLIFEM COMPLEX 



465 



In a revision of Asieropus from the Atlantic, 
Hajdu & van Soest (1992) described three 
species {A. hrasiliensis, A. vasiformis and A. 
niger). Asteropus radiocrusta differs from each 
of these in spiculation, at least by the absence of 
trichodragmata or the second size-class of 
oxyaster. This, in conjunction with other 
morphological and wide biogeographic 
differences, indicates significant variation at the 
species level. 

Stelletta Schmidt, 1862 

Kfyriastra Sollas, 1886: 187 (Type species: Myriastra subtilLs 

Sollas, 1886, by subsequent designation, see Sollas. 1888). 
Pilochrota Sollas. 1886: 189 (Type species: Pilochrota 

haekeli Sollas, 1886, by subsequent designation, sec de 

Laubenfels, 1936). 
Anthastra'SiOW-iX'A, 1886: 191 (Ty\iQ Anthastra pulchra 

Sollas, 1 886, by subsequent designation, see Sollas, 1 888). 
Dorypleres Sollas, 1888: 426 (Type species: Dory'pleres 

dendyi Sollas, 1888, by monotypy,). 
Incertae sedis: Astroplakina Dendy & Burton, 1926: 230 

(Type species: Astroplakina stelligera Dendy &. Burton. 

1926, by monotypy). 
Incertae sedis: Zaplathea de Laubenfels, 1950 (Type species: 

Zaplathea digonoxea de Laubenfels, 1950, by original 

designation). 

TYPE SPECIES. Stelletta gi-iihii Schmidt 1862, by 
subsequent designation (see Burton & Rao, 1932: 310). 

DEFINITION. Ancorinidae with fine-centrum 
euasters (oxyasters, strongylasters or tylasters) 
only as microscleres. 

REMARKS. Lendenfeld (1903) synonymised 
Myriastra^ Pilochrota and Anthastra with 
Stelletta, disregarding the presence of a second 
category of aster in the latter. However, Dendy 
(1916) found it convenient to retain Myriastra, 
but agreed with merging Pilochrota into it, since 
both had only one category of aster. Similarly, de 
Laubenfels (1936) maintained Myriastra as 
separate from Stelletta, but Bergquist (1968) 
again ratified the synonymy of Myriastra with 
Stelletta after assessing that three specimens in 
her collection were S. crater Dendy, 1924 which 
possessed two mutually exclusive categories of 
asters. 

Dorypleres has classically been considered to 
be closely related to Jaspis. and hence has been 
commonly placed in the Coppatiidae (or one of 
its junior synonyms), as either a distinct genus or 
a junior synonym of Jaspis. The genus was 
erected originally by Sollas (1888), without a 
generic diagnosis, but with a designated type 
species {Dorypleres dendyi Sollas, 1888), 
described as having two categories of aster. It was 
referred to Jaspis by Topsent (1904), although 



Burton & Rao (1932) noted that it did not 
conform to the typical structure of Jaspis 'having 
large oxeas only irregularly arranged, and asters 
of two kinds'. Indeed, Burton & Rao (1932) 
remarked how similar J. dendyi was to certain 
species of Stelletta, and were it not for the 
absence of triaenes, they claimed that they would 
have had little reservation in assigning the 
species to Stelletta. De Laubenfels (1954:^228) 
reversed Topsent's (1904) decision, restoring 
Dorypleres to generic status, defining the genus 
to include 'those species which have two or more 
distinct categories of asters, v^hQYQ Jaspis has just 
one category of aster'. This decision was sub- 
sequently reversed by Bergquist (1968: 33), 
noting that 'two categories of asters are not 
recognisable in sponges assigned to Jaspis 
dendyi\ a point corroborated by the present author 
after examining a slide of type material (holotype 
BMNH1889. 1.1.100). Consequently, in 
agreement with Bergquist's (1968) remarks, 
Dorypleres cannot be reinstated, as de 
Laubenfels (1954: 228) suggested, for 'those 
species (of Coppatiidae) which have two or more 
distinct categories of aster', and in which he 
placed Dorypleres splendens de Laubenfels, 
1954. 

Hajdu & van Soest (1992) briefly discuss 
Dorypleres, highlighting the differences between 
it and Jaspis and remarking on its Stelletta-\\kc 
nature (despite its lack of triaenes). They 
proposed that if the lack of triaenes was found to 
be a synapomorphic character, then Dotypleres 
may be reinstated as a JaspisAxko. Stellettid 
lacking triaenes. 

Recent morphological and chemical studies on 
jasplakinolide-containing sponges by Sanders et 
al. (1999) concluded in ratifying the synonymy 
between Dorypleres and Jaspis. They imdertook 
morphological studies at the supraspecific level 
on several nominal Jaspis species (including 
Dotypleres splendens de Laubenfels, 1 954), con- 
cluding that only one genus (Jaspis) was valid. 
This conclusion was based largely on similarities 
in skeletal composition and arrangement, the 
authors claiming that 'all species possess oxeas, a 
confused choanosomal arrangement and para- 
tangential arrangement of small spicules at the 
surface' (Sanders et a!., 1999: 526). Although 
strictly correct, they did not distinguish between 
the nature of the 'small spicules' comprising the 
ectosome of each species, Dorypleres splendens 
has an ectosome containing a thin crust of 
oxyasters, whereas Jaspis, including the type 
species, J. johnstonii (Schmidt, 1862), has a 



466 



MEMOIR.S OK T I It QUEENSLAND MUSEUM 



tanger.uat cctosatne c<>nl|>a86d Af OKeote 
5piQult;&. in fi«?iitrji6t lu ihcir suprnspccifiB 
iE^lQ)dIu5ticMi5'l«fl&d cm morphological comparisons, 
their eoncltisions based on chemical analysCvS 
■pertain only to the species level of classification, 
<int) hence iwc unanlc io pro\idc any reliable 
giiru:ric charactcrislic. Saudcrs et ai. ( 1999) con- 
cluded Ihai 'all of ihojusplakinolidc-containing 
sponges studied were found to be conspccitic' 
and that \Iuspl^ splenJam {dcLmha\(^h, 1^54) 
IS the scmor-mosl iivaikible name for these 
specimens'. A consequence rrf this lo-jic is that 
Ihc present chetnical naini ;.i-.pl,ikuioh(Je is 
misleading since it pertains only to a single 
Species ('Viffjpp^gm splemlem')' Hn6 n6l to a 
genus (liaLUiostfelllwICPS 2''*'^ perhaps should he 
fLMuimcii nsinjt Sll!hC-<teri^''"'^'" Npci ies 

name s/^^t m?t7;5 * (pcijiups 'splcndcnctlkle') 
(alliiougji nom^^ndaWalty this probably not u 
tcasibic propositidny. 

■^ ,1.1 ! •i.-n.f.ih'n'.s Joes ium ;i lainiential 

cciosumc at oxcotc spicules, it lalls outside the 
dtaij;nosb of Copp^idfte. ftild is COM^idi^We 
to be n Stelletta lackmjz triaencs.. 

SaDders er ^. (1999) iiUa ftyttottj/tnised- die 
itionotypic genus Zaplefheo'ie Laubenfeti^ f*>50 

willi ./,/.\//;v. i>;ised oi) celosonial skeletal 
arrangcTncTir ol die type species (/. c/o^^onc'X.- a de 
Laubentcls, l^).s()), again w iiiu>ui . lueidaiinu die 
cvtfiponenl spicule types, their lUusirulion ol a 
cross section of the type spcdnttaicctosomc docs 
not show the diagnostic tangential layer of 
oxeotes Uiat is typical of Coppatiidae, and the 
un,uinal desenpiion ( dc Laubeniels, 1950) suites 
that there is no sharply defined cortical re^jiop'. 
The iwiee henl 'nucroseas', described-by de 
Laubenivls as being diagni^stic for the genu$> 
'wfcre inlerpreied by Sandarel (1999) as being 
a diagnostic feature at the species level only. It is 
H|ieeulated ihiU Znplaihea is also a Si*.'lli fia 
laekinj: inaencs. Donplcres ^b\^\c, although 
lurther tescareh is needed to eunllrn! (his. 

Astroplakina Dendy & Burton, i 92(>, is a mono- 
typic geruis eomaining onlyvt. t^^lUgera Deiidy 
& BurlotL i 926. The type species Is described as 
fiaving a range of spicule types, ranging from 
diaels to otMacts. It is apparent Irom the ongmal 
figures that the diacts arc oxeotes and the 'triads 
to octacts' are oxyasters. Indeed, nend> & 
Uurlon (1926) noted \h<t smidanlies ol these 
.spicules to the oxy- or sliongylasicrs of the 
Steiidtidae {- Ancorinidac>« particularly w here 
Ihe Are microspincd. Unlortunaieiy, the 
skeletal stnictancwas noi desctifeed b^e^uso^the 



specimen was dry and unable to be sectioned 
jufeqsate^. «A£ fw Zciphth^^ sod' DoryptenffSs 
Mirr^p^aktm fipecnlated to ft Bfefletta 

Uickint: triaenes, alfhougli fiitthtStiSOnfuTnaiion is 
ncccssaiy, paiiicuiarly iclatitB^td'&e cctosomal 

S(«iretttt|tbefXulala (Carler. |8B6) 

(Rgs 1, 6, Tabic 6) 

MATERIAL. KrCTOTYPf-: nMNHI«W.I2.15.434 
(dry): POit Phillip Heads. Vlcioria OTHER MATERLAL: 
BMNM 1 954.2. 1 2.251; maAed *!Slide of type' (prepiuwi by 
A. bendy). 

HABITAT DISTRIBUTION. 3-6m depth; on 
gninite boulders with algae; Victoria, Bass Strait. 

DrACfftoSlS. Massive, with irregular ttfbeincics, 

ridij;es and lobcS}* dull -purple lo red-brown alive; 
numerous itiirtutc csculcs in depressions between 
Uihercles; dislinclly arenaceous surface; COrticfll 
ectosome ot;sand-grains largely obscurliigniinutie 

posed of oxcas in confused arrangement largely 
obscuring scattered oxyasiers; megascleres 
flcxuous; oxcas in two si/e classes diflerentiatcd 
mainly by their thickness (thicker oxeas length 
63(M78S)-863, width ll-(lt.)-i9: thinner oxcas 
lcngth3 U-(52|)^84, width 2-(5)-8)i n««|ftloA 
plagioiriaene^ m tWo sixe classes (thicker triaenes 
rhabd length h'^O-il'l 1 )-82 ! . clad leneth 1 0-( 1 ^i)- 20; 
thinner triaenes rhabd length 535-(6 1 1 clad 
length 7-(7,5).8)iiisicfS«pjlWd i»cy««e» 
tl0)-12pm). 

DFSCF<IPTIQN. ^:^^ipt'. Massive, covtred witJi 
iMisied. irrcgutar nbdulosc ridges, lobes and 
tubercles OH top and sides, with furrows and 
dcpressioni^ b^Vi/een; tubercles 3-l2mni thick 
and 5-lgmm di^: Heiglit l.leWf width 5.S X 
12cm, 

CohuK Dull pvnple-bpown (Cartn; t886a) W 

rcddish-hrown (Wiedenmayer, 1 989);!iVij(at 
fresh; dry leclotype with greyish beigc^biow^. 
cortical ccloson-ic (Munseli 2.5Y Mi 
liglit beige cljoaiiosome (2,5V 8/3). 

Osi'u(09, jftmeMss, almost iiidistfnci oscules, 

approximately 0.5mm diameter, appear as 
slightly darker spots in the ftiirows and sulci 
between tubercidaC&SDbes. 
Texfure. Dr>' leclotype is hard, stony* coaree; wcl 
niaierial is reported as fuTn, barely compressible* 
. ^ily torn (\Medemnay^» 1989). 



RESOLVING TmJASPlSSTELUfEHi COMPLEX 



467 




FIG 6 SteIleitaiubereUlm(CBTi&, 1886) (lectotvpe BMNH]8fi6J;2.15.434). A, lectotype; B, seel ion Uirough 
peripheral skeleton (BMNH1954.2.12.253); C, thick oxea; D, thin oxea; E, thick ortho/pla^trUeQe; f,6 
cladomes of thick ortho/piagiotriaenes; H, thin ortho/plagiottiaene; I, thin ortho/plagiotria»ve claAjriie; 
oxyastet. 



Surface charaet^^Mitt Highly tuberculate and minute oxy asters are scattered tihrQi^Crut but are 

conspdcuously arenaceous. largely ^(bseurcd by the sand. 

Ecio.some. Approximately 1 -2mm thick.foyming Chof^S^- Sketeton comprised e^acojifusioix 

a cortex that is distinct from choanost?m^ dtietq ofsittgleaiidVeiy loose paqcispicutatbOT 

the pfesence of abundant Iwge san^-grains; oxeasthatfafgely obscure the abundant Qxyasl^ 



468 



MEMOIRS OF THE QUEENSLAND MUSEUM 



TABLE 6. Comparison between present and published descriptions of Stelletta tuberculata. Measurements in 
|im, denoted as range (and mean) (N=25 in present study except where noted). L = length; W = width; RL = 
rhabd length; CL = clad length; D = diameter. 





Oxeas (Thick) 


Oxeas (Thin) 


Triaenes (Thick) 


Triaenes (Thin) 


Oxyasters 


Lectotype 

BMNH1886.12.15.434 
Present study 


L 630-(788)-863 
W n-(I6)-19 


L3l3-(52I)-684 
W2-(5)-8 


RL670-(771)-82I 
CL 10-(16)-20; (N=7) 


RL 535-(6n)-668 
CL 7-(7.5)-8; (N-4) 


D6-(10>]2 


Syntype, original descrip- 
tion (Carter, 1886a) 


L762 
W 13 


Not described 


Not described 


Not described 


D8.5 



in the dense matrix; ortho/plagiotriaenes present, 
with dads just below the cortical ectosome and 
rhabds directed vaguely inward. 

Megascleres. (Refer to Table 6 for spicule 
dimensions) Oxeas in two size categories dit- 
ferentiated primarily by their thickness and 
extent of tlexion; thicker oxeas lightly flexuous, 
infrequently fusiform, with acerate to finely tele- 
scoped ends. 

Thinner oxeas similar in geometry but much 
more flexuous. 

Ortho/plagiotriaenes in two size classes also 
differentiated by their thickness; rhabd tennin- 
ations slightly telescoped; clads stumpy. 

Microscleres. (Refer to Table 6 for spicule di- 
mensions) Oxyasters with 10-17 lightly tapering 
rays that have 10-20 recurved spines on distal 
half, thus giving a slightly tylote appearance 
particularly when examined using light micro- 
scopy; centrum approximately 25% of spicule 
diameter, 

REMARKS. The original description by Carter 
(1886a) was obviously based on specimen 
BMNH1886,12.I5.434, nominated here as the 
lectotype, since the other two syntypes are clearly 
different species. One syntype (BMNHI886. 
12. 1 5.113) is a Crella of uncertain specific identity, 
and at first was thought to have been a mislabelled 
holotype of Carter's ( 1 885) Echinonema (Crella) 
incnistans (BMNH1886. 12.15.123) (i.e. 
considering the similar registration numbers). 
However, this is not the case because this latter 
specimen was also examined by the author and is 
different again. The remaining syntype (BMNH 
1886.12.15.146) is clearly yet another species 
(described above as A. radiocmsta sp. nov.). 

Stelletta titberculata is reinstated here as a valid 
species, distinct from J. stellifeni in several 
important respects. Stelletta tuberculata has both 
oxeas and triaenes as megascleres (cf ./. stellifera 
which only has oxeas). Further, S. tuberculata has 
a highly arenaceous cortical ectosome of sand- 
grains, largely obscuring scattered minute oxy- 
asters (cf a tangential layer of oxeas in confrised 



arrangement). The vestigial nature of the triaene 
clads makes it difficult to properly resolve their 
form as orthotriaene or plagiotriaene. 

As Wiedenmayer (1989) remarked, there are 
very few Stelletta species that contain foreign 
detritus and relatively rare, reduced triaenes. He 
addressed the differences between these species 
in his remarks for S. arenitecta, which is syn- 
onymised here with S. tuberculata, 

Rhabdastrella Thiele, 1903 

Aurora Sollas, 1888: cxxxix, 187 (preoccupied, junior 
homonym of Aurora Ragonot. 1887 (Lepidoplcra)) (Type 
species: Stelletta globostellata Carter, ! 883, by original 
designation). 

Rhabdastrella'X\\\Q\Q, 1903: 934; Bergquist, 1968: 54. 
Diastra Row, I9U: 300; Bergquisi. 1968: 54 (Type species: 

Diastra sterrastraea Row, 191 1, by monot>py). 
Aurorella De LaubenfeLs, 1957: 245 (nomen novum fox Au- 

rara Sollas, 1888); Wiedenmayer, 1989: 21. 

TYPE SPECIES. Coppatias distincnis Thiele, 1900, by 
original designation. 

DEFINITION. Ancorinidae with thick centred 
euasters (oxyspherasters or spherasters) in a 
cortical ectosome. 

REMARKS. Aurora was originally proposed for 
Carter's Stelletta globostellata and S. reticulata 
because they possessed large oxyspherasters. 
Lendenfeld (1903) merged the genus with 
Stelletta, followed by Hentschel (1909). Dendy 
(1916) argued that it was desirable to retain 
Sollas 's genus, since the large (oxy)spherasters 
form such a characteristic and well-detlned 
feature, and are known from many species. He 
also proposed the addition of Diastra sterrastrea 
Row, 1911, A. cribroporosa Dendy, 1916 and 
Coppatias (Rhabdastrella) distiuctus Thiele, 
1900, Thus, he proposed the synonymy of 
Rhabdastrella and Aurora. He also suggested 
that the type species o{ Aurora, A. globostellata, 
did not have trichodragmata, as Sollas 1888 sug- 
gested (corroborated in the present study). Dendy 
also asserted that the loss of triaene megascleres 
has taken place several times within Aurora^ 



RESOLVING THE JASPIS STELLIFERA COMPLEX 



469 



giving several examples of similar species 
differing in the presence of traienes. 

De Laubenfels (1957) proposed the name 
Aurorella to replace the preoccupied Aurora, but 
still maintained it as distinct from RhabdastJ-ella 
(which he merged into Dorypleres). He restored 
Diasira to frill and valid generic status based on 
the possession of sterrasters. 

Bergquist (1968) synonymised Aurora and 
Diastra into Rhabdastrella on the basis of 
Dendy's (1916) argument. She used Rhabd- 
astrella to receive d\\ Aurora species because the 
latter name was pre-occupied, also drawing 
attention to Dendy's observation that tliree pairs 
of species within Rhabdastrella (s.s), Diastra and 
Aurora were only distinguishable by the presence 
or absence of triaenes. It appears that these 
observations have led to the modification of the 
definition of Rhabdastrella (e.g. Wiedenmayer, 
1989: 21) to include the character 'with reduced 
triaenes or without triaenes . . .'). 

Hechtel (1 983) mQ& Aurorella as a subgenus of 
Rhabdastrella on the basis that it lacked triaenes. 
As has been suggested (Hajdu& van Soest, 1992) 
for the Ancorinidae (and Coppatiidae), the lack 
of triaenes is a suspect diagnostic character. 
Evidence is given here to support this, whereby 
R. globostellata is shown to have a gradation of 
triaene development, ranging from well- 
developed, through to vestigial or absent. 

Rhabdastrella globostellata (Carter, 1883) 
(Figs 1,7, 8C-F. Table 7) 

SteUetta globostellata Carter, 1883: 353-354. 
Aurora globostellata 1888: 187-188. 

Stelleltmopsis carteri Ridley, 1884: 476. 
Coppatias carteri SoUas, 1888: 208-209. 
Jaspis stellifera BQTgquist, 1969; 69. 

MATERIAL. HOLOTYPE: BMNHl 883.5.3. 1 (dry): 
Galle, Sri Lanka (Ceylon), coll. Dr. Ondaatji. HOLOTYPE 
of Stellettinopsis carteri Ridley, 1884 BMNHl 882. 
2.23.276 (wet): Prince of Wales Channel, Torres Strait, 
coll. R.W. Coppinger. OTHER MATERIAL: Australia - 
Western Australia, NTMZ3352, QMG301116, 
QMG301 142; Northern Territory, NTMZ96, Z582, Z588, 
Z599, Z1325, Z2182, Z3248, OMG303634, G313548; 
Queensland, NTMZ4011, QMG300041, G30n79, 
G303170, G303487, G303509, G304341, G304450, 
G304606, G304481, G304884, G305457, G305779, 
G306240, G313432. G313472, G313508, G313589, 
G314452, G314563. G314624, G315114, G315227, 
G315249, G31 5503. New South Wales, QMG30 1398. Fiji - 
QMG3 12735 (NCIOCDN-4165-M), QMG3 12803. 
Malaysia - OMG301224, G301227, G301228, G304613. 
Palau - QMG305951. Philippines - QMG3 12576. Tonaa - 
QMG313264. Vanuatu - QMG306826. G306893 
(ORSTOM R1624). 



HABITAT DISTRIBUTION. Intertidal-53m 
depth; on coral reef, rocky outcrops, broken reefs, 
reef flats, drop-ofYs and overhangs; tropical and 
subtropical Indian and Pacific Oceans; Sri Lanka, 
Singapore, Zanzibar, Indonesia, Malaysia, Palau, 
Philippines, Vanuatu, Fiji, Tonga, Australia; N 
Western Austi^alia, Northern Territory, Queens- 
land, Great Barrier Reef, N New South Wales 
coast. 

DIAGNOSIS. Massive, globular, subspherical; 
brown to yellowish-tan cortex, with yellow 
choanosome alive; apical depressions with 
numerous small oscules in clumps; ectosome of 
oxyspherasters; peripheral choanosomal skeleton 
variable, with plumose brushes of oxeas and 
orthotriaenes (sometimes absent) thai have 
rhabds directed inward; deeper choanosome 
contains oxeas in confused arrangement, with 
oxyasters in variable abundance between spec- 
imens; oxeas (length 220-(8l4)-1521, width 
0.5-(13)-38), orthotriaenes (rhabd length 
70-(600)-1309, clad length 7-(104)-239), oxy- 
spherasters (diameter 5-(36)-91), oxyasters 
(diameter 3-(28)-96). 

DESCRIPTION. Shape. Dry holotype is incom- 
plete, amorphous, and has an irregularly folded 
surface which Carter (1883: 353) described as 
'corrugated'. Living and wet-preserved spec- 
imens are typically globular, subspherical, 
sometimes raised on a short, thick base that is 
attached at several points, commonly with one to 
several shallow, concave depressions (up to 5mm 
depth) on apical surface. Incomplete holotype 
5.5cm long, 3.5 x 2.5cm wide; larger specimens 
up to 30cm high, 45 X 45cm wide. 

Colour Dry holotype has dull greyish beige- 
brown (Munsell 7.5\1l 7/2-6/4) ectosome, with 
beige-tan choanosome (7.5YR 6/4). Living 
specimens have variable ectosomal colouration 
between specimens, ranging from from 
yellowish-tan (2.5Y 7/4) to deep chocolate brown 
(7.5YR 4/4), and occasionally dusty purple 
(10R7/2), how'ever the choanosome is always 
vivid yellow (2.5Y8/10). 

Oscules, Not visible in incomplete, dry holotype, 
but Carter (1883: 353) originally describes 'vents 
congregated in one part of the surface'. Recently 
collected material typically has numerous (about 
20-40) small oscules ( 1 -4mm diameter ) clustered 
in one to tliree, large concave depressions (up to 
5cm deep) on apical surface, with larger oscules 
in more central regions of clusters. 



470 



MEMOIRS OF THE QUEENSLAND MUSEUM 




FIG 7, Rhabda^itrella ^ob^stelkitu (Carter. ! SS3). A. whole wel specimen from Vanuatu (QMG306893); B, 
holotype of Stel/eitimpsis cancn Kidle>, 1884 (BMNH1SS2.2.23.276 wet: Torres ^ajl, AuslialiaJ^ C, 
hok%pe(BNTNH1883.5.5.1 dn; Sn Lanka); D.l:. sections through peripheral skeletonsshowuigd^^ 
Structure between low latitude (D, specmiea QMO306S93: Vanuatu) and higher latitiMi& (]^/Ql(IQ3B4^^^ 
Heton lsland, GBR. Australia) material; F, F^-I,F, examples of spfoiilatJcmfliHbeH^JjettVeeft ip 
(F-J, holo^e of S, carterJK\^\ty, 1884 BMNH 1882.2,23 .276: Torres Strait, Australia; F'-T', pNi(33068^3: 
V«nuata);^#', oxeas; G,G% ortIiotriaenes;H,H', oxyasterS; l,r, oxyspherasters). 



RESOLVING THE JASPISSTELLIFERA COMPLEX 



47] 



TABLE 7. Comparison between holotype and other material of Rhahdastrella glohostellata by region. 
Measurements in )am, denoted as range (and mean). L = length; W ^ width; RL = rhabd length; CL = clad length; 
D = diameter. 



Region/material 


Oxeas 


Orthotriaenes 


Oxyspheraster euasters 


Oxyaster euasters 


HololypCi Sri L.anka, 
BMN}f!883.5.3.1 


T fi^H CHI 1 \ 1 nin 

L, OZo-^^o J 1 I' 1 \JD\) 

W5-(14)-23 


CL 65-( 126)- 172 


D8-(25)-55 


D ] U(25)-72 


Hololvpc of Stdl^ttinf^f^^i^ 
carltri. Torres Strait. Qld. 
BMNHIS82.'^ .23.276 


L 730-(859)-1020 
W I3-(18)-25 


r<.L 560-(647)-810 
CI. 70-(93)-l 10. (rare) 


D6-(42)-64 


D23-(36)-46 


Sabah State, Malaysia. 
4 specimens 


L 611-(1072)-1521 
W4-(21)-33 


RL 254-(835)-1309 
CL 21-(127)-239 


1 , 

D 15-(43)-61 


D8-(26)-56 


Philippines, ! specimen 


L790-(970)-l 150 
W6-(18)-26 


RL 326-(657)-946 
CL 37-(102)-138 


D 20-(46)-63 


D 33-{48)-63 


Palau, 1 specimen 


L566-{756)-963 
W2-(7)-l5 


RL 320-(559}-753 
CL50-(104)-166 


D8-(34)-50 


D6-(15)-21 


Vanuatu. 2 specimens 


L310-(641)-875 
W2-(5)-9 


RL 220-(43l)-638 
CL 7-(38)-85. (rare) 


D9-(27)-43 


D 7-(22)-39 


Fiji. ! specimen 


L 570-{736)-940 
W3-(10)-16 


RL 330-(356).382 
CL 43-(52)-65. (rare) 


D 15-(36)-54 


D 15-(18)-23 


Tonga. I specimen 


L599-(976)-ll6f; 
W 10-(20)-37 


RL 654-(726)-848 
CL 94-(148)-185 


D 11 -(40 1-66 


D9-(15)-23 


N Western Australia. 
3 specimens 


L472-(863 )-1425 
W4-(12)-19 


RL 219-(806)-ll07 
CL53-{128)-215 


Dn-(37)-72 


D 10-(43)-71 


Northern Territor>', Australia. 


L687-(998)-1456 
W4-(19)-38 


RL367-(818)-I240 
CL 53-(122)-215 


D 6-(42)-91 


D 11 -(47 1-96 


Queensland, Australia, 
13 specimens 


L220-(54!)-936 
W0.5-(6)-15 


RL70-(112)-247 CL55 
(Very rare: 2 specimens) 


D 5-(29)-55 


D 3-(22)-45 


N New South Wales, Australia, 
1 specimen 


L265-(527)-718 
W 2-(7)-i3 


None 


D 15-(36)-53 


D 14-(24)-31 



Texture. Dry holotype is hard; fresh and wet- 
preserved material is firm, compressible, and 
leathery. 

Surface characteristics. Opaque, optically smooth, 
uneven, with low, rounded tubercles, corrugat- 
ions, bumps and ridges fomiing a tuberculate 
surface becoming smoother toward the base; 
extent of tuberculation varies between spec- 
imens, ranging from prominently tuberculate to 
nearly completely smooth. 
Ectosome. About 150-500 thick; skeleton com- 
posed exclusively of oxyspherasters. Variable 
degrees of packing of oxyspherasters occurs, 
ranging from very densely packed in tropica! 
material, to relatively sparsely packed in 
subtropical specimens. Canals, approximately 70 
diameter, regularly traverse the ectosome. Sub- 
ectosomal region is relatively clear of microscleres. 
Clwanosonie. Deeper choanosomal skeleton 
consists of a confusion of loosely scattered single 
oxeas and paucispicular bimdles of oxeas. These 
bundles become more ordered in the peripheral 
choanosome, where they may also include ortho- 
triaenes, fonning variably distinct paucispicular 
plumose brushes, with the rhabds of the triaenes 
directed inward and clads supporting the ectosome. 
There is substantia! variability in absolute 



abundance of microscleres and relative 
abundance of microsclere categories between 
specimens, without any obvious correlation 
between geographic regions or latitudinal 
gradients. 

Megascleres. (Refer to Table 7 for spicule 
dimensions) Oxeas, slightly bent, with finely 
telescoped ends; variations very rare but include 
styloid forms and oxeas with temiinations that 
are split or sharply bent (similar to promonaenes). 

Orthotriaenes, with variable clad development, 
with rhabds tapering toward frisiform, hastate or 
faintly blunt terminations; each cladome contains 
three identical clads with shapes ranging from 
typical geometries, to lightly telescoped, sharply 
angled near tips, or stunted fonns. Orthotriaenes 
more commonly observed and robust in tropical 
rather than subtropical specimens where they 
may be highly vestigial or apparently absent. 

Microscleres. (Refer to Table 7 for spicule 
dimensions) Oxyspheraster euasters, with 9-18 
conical rays; centruin approximately 29% of 
spicule diameter; rays may be entirely smooth or 
have up to 15 microspines on distal end. 

Oxyaster euasters, with about 9-17 lightly 
tapering to isodiametric rays, with approximately 
11-25 recur\ ed microspines on the distal half. 



472 



MEMOIRS OF THE QUEENSLAND MUSEUM 



thus giving a slightly tylote appearance under 
Hght microscopy; centrum approximately 1 0% of 
spicule diameter; variations ver}' rare, with only 
one spicule displaying forward projecting 
microspination over entire ray length. 

REMARKS. This is undoubtedly the same 
species referred to by Bergquist (1969) as 'J. 
stellifera from Heron Island, GBR, with the 
qualification that her specimens are represent- 
ative of subtropical material found typically to 
have reduced spicuiation. In agreement with her 
published remarks, and corroborated by more 
recent surveys of this region, it is one of the most 
common and more prominent species of sponges 
on the reef Hat, easily recognisable for its 
massive, subspherical shape, brown exterior and 
distinctive (mango-like) bright-yellow interior. 
Spicule diversity and size are similar between 
Bergquist's (1969) and recent collections, with 
the qualitlcation that the tylasters described by 
Bergquist are actually oxyasters with micropines 
clumped near the terminations and only seen 
properly under SEM. This species, however, is 
clearly different from J, stellifera in many 
significant respects. 

While both species are essentially subspher- 
ical, they diOer greatly in size and colouration, 
but most significantly in ectosomal skeletal 
structure and spicule compliment. Jaspis stel- 
lifera is about 4.7cm in largest dimension and 
pinkish-white to purple, whereas R. ^lobostellafa 
is at least up to 45cm in diameter and has a brown 
exterior and bright yellow interior. Both species 
have choanosomal skeletons that are essentially 
confused arrangements of oxeas, with oxyasters 
scattered in the interstices. However, the ecto- 
some of R. ^lohosteilata has a distinct layer of 
oxyspherasters, in contrast to that of of J. stel- 
lifera. which is comprised of a tangential layer of 
oxeas. Further, ./. stellifera has only oxeas and 
oxyasters as spicules, whereas R. globostellata 
also has orthotriaenes and oxyspherasters. 

The synonymy of Stellettinopsis carteri 
Ridley, 1 884 with R. globostellata is based on the 
presence of orthotriaenes and oxyspherasters in 
the holotype of .S'. carteri in addition to the oxeas 
and oxyasters as originally described. Further, 
spicule sizes (refer Table 7) he within the range 
described for R. globostellata. Unfortunately, the 
type specimen is no longer complete, as origin- 
ally described by Ridley (1884), now consisting 
only of a small fragment (height 8mm, width 
21 X 25mm), w ith only a small portion of surface 
intact. However, the original description of the 



gross-morphology of 5. carteri is consistent with 
characteristics of R. globostellata (e.g. having a 
'short cylindrical stalk passing gradually into a 
massive, somewhat flattened upper portion*; 
colouration being tan with a yellow interior; and 
with an undulate, dimpled, corrugated surface). 
Unfortunately, the ectosome is barely intact in the 
holotype of S. carteri, and consequently it is 
difficult to ascertain its true nature. It is vaguely 
distinct from the choanosome (although not 
explicit in Ridley's (1884) original description), 
being smooth and probably composed of oxy- 
spherasters. This synonymy is further supported 
by the type locality of S. carteri being well within 
the distribution of i?. globostellata. 

In R. globostellata, orthotriaenes were more 
abundant in specimens from more northerly 
tropical localities, where spicuiation was 
generally more highly silicitied. By comparison, 
specimens from more southern regions (central 
GBR to N NSW), typically lacked iriaenes and 
spicules were more poorly silicified and less 
robust. Tliis is similar to the latitudinal trend 
observed by Hooper & Bergquist (1992) for 
Cynibastella (Axinellidae), and Hooper (1996) 
for Clathria (Thalysias) vulpina. Failure to 
recognise the absence of triaenes in specimens 
may perpetuate misidenlifications of this species 
in the fliture, and so the distinctive shape and 
colouration ofthis species will undoubtedly remain 
an important, albeit superficial, distinguishing 
features. 

Tlie reassignment of tropical sponges, previously 
misidentified as V. stellifera\ to Rhabdastrella 
globostellata is based primarily on morphologi- 
cal evidence and is supported by chemical 
evidence in the possession of malabaricane-type 
triterpenes (e.g. Ravi et al., 1981; Ravi & Wells, 
1982), determined by van Soest & Braekman 
( 1 999) to be a good chemotaxonomic marker for 
Stellettids. It is possible that V. stellifera^ from 
Japanese waters are also misidentified specimens 
of S. globostellata since they too have been reported 
to contain malabaricane-type triterpenes (Tsuda et 
al., 1991; Kobayashi etal., 1996). 

DISCUSSION 

Examination of all type material previously 
assigned to, or associated with, "Jaspis stellifera^ 
revealed that many important details were 
omitted from original descriptions. Failure to 
recognise these details has certainly contributed 
to an oversimplified synonymy for this species. 
For example, re-examination of type material 
showed clearly that the syntype series of 



RESOLVl>jG JllEJASPISSTELLIFERA COMPLEX 



473 




FIG. S- A-B, ././Y''^ , ri^fucnrrui^cnus sp. nov. (OMG3I2071, Port Davey, Tasimania, lOm, photo NCf); A, 
specimen in situ; R. close-up \ ie\\ of oscular ridge surface. C-T. R/iiihJcisDT/la silohostelhta (Carter. 1 R83); C, 
N rMZ2 1 82. Darwin, N i; iniertitlal,photo J. I (ooper: D, QMG306jS^3. Emae, Vanuaiu, 20m, photo ORSTOM; 
i:, Q\\g:> t >432. hteron Isiani QI<1, intettklttU photp authBr^ F, Q\4G3048S4, Mudjimba Island.. QW. I2ni* 
pholo J. Hooper. 



474 



MEMOIRS OF THE QUEENSLAND MUSEUM 



f SteUeifinopsis tuherculata was composite, 
containing specimens from difTerent orders. This 
example is perplexing because the original 
synonymy was proposed by Shaw (1927), under 
the direct supervision of Maurice Burton at the 
BMNH, who had complete access to the vast type 
collections housed there. It is probable, therefore, 
that this synonymy was based on superficial 
comparison of type material rather than re- 
examination of histological preparations. 

In recognising the oversimplification of Shaw's 
(1927) synonymy, Bergquist ( 1969) reinstated J. 
coriacea and ./ purpurea as \'alid species, based 
on two supposed inconsistencies in the published 
literature, both of which were demonstrated here 
to be invalid. In contrast, the proposed synonymy 
of ./. slellifercu based on redescribed type 
material, incorporates only ./ coriacea and J. 
purpurea. Consequently, J. stellifera is not as 
widely distributed as reported by Hooper & 
Wiedenmayer (1994). Its corroborated 
distribution appears to be restricted to waters 
between Victoria and Tasmania. Most of the 
remaining species previously placed in the V. 
stellifera' complex also appear to have very 
limited distributions, with the exception of R. 
globosteliata which has an apparent widespread 
tropical/subtropical Indo-Pacific distribution. 

The clue to misidentified tropical ^md sub- 
tropical populations of R. globosteliata as "J. 
stellifera'' is largely based on the presence of 
triaenes within specimens, even though these 
range from present to vestigial or sometimes 
absent in individual specimens. It was only 
through thorough examination of many specimens 
that this anomoly concerning presence/absence 
of triaenes was recognised. Triaenes were more 
common in tropical specimens, where spicules 
were generally more robust than in southern pop- 
ulations. The apparent lack of triaenes in some 
material, or failure to recognise their vestigial 
occurrence in other specimens, may lead easily to 
the misidentillcation of this species as Jaspis 
(which by definition lacks triaenes). Tlie graded 
development of triaenes across tropical to 
subtropical waters has implications regarding the 
debate surrounding the relationship between 
Coppatiidae and Ancorinidae. According to 
present diagnoses, the presence of triaenes 
confirms that this species belongs to Ancorin- 
idae, whereas specimens lacking triaenes could 
be justifiably included in Coppatiidae. The rare, 
vestigial, or complete loss of triaenes in R. 
globosteliata provides evidence supporting the 
proposition of Hajdu & van Soest (1992) that the 



absence of triaenes is a dubious synapomorphic 
character used to separate Coppatiidae and 
Ancorinidae. 

ACKNOWLEDGEMENTS 

I thank the following for their assistance with 
this study: Clare Valentine (BMNH, London), 
Ruth Desqueyroux-Faunde/ (MNHN, Geneva), 
NTM, Darwin, for loan of specimens and tvpe 
material; Pat and Lori Colin, CRRF Patau for 
material; John Hooper for collections from 
throughout tropical Australia, Fiji, Malaysia, 
Philippines, Tonga and Vanuatu; the Queensland 
Museum lor facilities, field support, and 
materials; two anonymous reviewers for their 
valuable comments on the manuscript; and John 
Hooper for his mentoring, taxonomic guidance 
and valuable comments regarding this paper. 

LITERATURE CITED 

BERGQUIST, P.R. 1965. The sponges of Micronesia, 
part 1 . The Palau Archipelago. Pacific Science 1 9: 
123-204. 

1968. The marine fauna of New Zealand: Porifera, 
Demospongiae, Part 1 (Tetractinomorpha and 
Lithistida). New Zealand Department of 
Scientific and Industrial Research Bulletin 188: 
1-105. 

1969. Shallow water sponges from Heron Island. 

LIniversit\' of Queensland Papers, Great Barrier 
Reef Committee, Heron Island Research Station 
1(4): 63-72. 

BOURY-ESNAIIIT, N. & RUTZLER, K. (eds). 1997. 

Thesaurus of sponge morphology. Smithsonian 

Contributions to Zoology 596. 
BURTON, M. 1934. Sponges.'Scicntific Reports of the 

Great Barrier Reef Expedition 1928-29. 4(14) 

5L3-621 (British Museum (Natural History): 

London). 

BliRTON, M. & RAO, M.S. 1932. Report on the 
shallow-water marine sponges in the collection of 
the Indian Museum. Records of the Indian 
Museum 34(3): 299-356. 
CARTER, H..1. 1 875. Note introductor\ to the study and 
classification of the Spongida. Annals and 
Magazine of Natural History (4)16: 1-40, 
126-145, 177-200. 

1879. Contributions to our knowledge of the 
Spongida. Annals and Magazine of Natural 
Hislor\- (5)3: 284-304. 343-360. 

1883. Contibutions to our Knowledge of the 
Spongida - Pachytragida. Annals and Magazine 
of Natural Histoty (5)11 : 344-369. 

1885. De.scription of sponges from the neighbour- 
hood of Port Phillip Heads. South Australia. 
Annals and Magazine of Natural History (5)16: 
347-368. 

1886a. Description of sponges from the neighbour- 
hood of Port Phillip Heads, South Australia. 



RESOLVING THE JASPIS STELUFEM COMPLEX 



475 



Annals and Magazine ofNatural llistor>' (5)18: 
126-149. 

1886b. Supplement to the descriptions of Mr. J. 
Bracebridge Wilson's Australian sponges. 
Annals and Magazine ofNatural History (5)18: 
445-466. 

DENDY, A. 1905. Report on the sponges collected by 
Professor Herdman, at Ceylon, in 1902. Pp. 
57-246. In Herdman, W.A. Report to the Govern- 
ment of Ceylon on the peral oyster Fisheries of the 
Gulf of Manaar. 3(18): 57-246 (Royal Society: 
London). 

1916. Report on the Homosclerophora and Astro- 
tetraxonida collected by H.M.S. 'Sealark' in the 
Indian Ocean. In Reports of the Percy Sladen 
Trust Expedition to the Indian Ocean in 1905, 
Volume 6. Transactions of the Linnean Society of 
London, Zoology 17: 225-271. 
1924. Porifera. Part L Non- Antarctic sponges. British 
Antarctic ('Terra Nova') Expedition, 1910. 
Natural history report. 6{3): 269-392. (British 
Museum (Natural Hislon ), Zoology: London). 

DENDY, A. & BURTON, M. 'l926. Records on some 
deep-sea sponges from the Indian Museum 
collected by the R.I. M.S. Investigator'. Part 1. 
Hexactinellida and Tetraxonida (pars.). Records 
of the hidian Museum 28: 225-248. 

FUERST J.A., WEBB, R.I., GARSON, M.J., HARDY, 
L. & REISWIQ H.M. 1999. Membrane-bound 
nuclear bodies in a divers range of microbial 
symbionts of Great Barrier Reef sponges. Mem- 
oirs of the Queensland Museum 44: 1 93-203. 

FUSETANI, N. & MATSUNAGA, S. 1993. Bioactive 
sponge peptides. Chemical Review 93: 1 793- 1 806. 

GRAY, J.E. 1 858. Description of a new genus of sponge 
(Xenospongia) from Torres Strait. Journal of 
Zoology 26: 229-230. 
1867. Notes on the arrangement of sponges, with 
the description of some new genera. Proceedings 
of the Zoological Society of London 1 867: 492-558. 

HADJI], E. & SOEST, R.W.M. van 1 992. A revision of 
M]mX\c Asteropiis Sollas, 1888 (Demospongiae), 
including a description of three new species, and a 
review of the family Coppatiidae Topsent, 1898. 
Bijdragen lot der Dierkunde 62(1): 3-19. 

HECHTEL, G 1983. New species of marine Demo- 
sponges from Brazil. Iheringia. Serie Zoolosica, 
Porto Alegre 63: 59-89. 

HENTSCHEL, E. 1909. Tetraxonida. 1. Teil. 2(21): 
347-402. In Michaelsen, W. & Hartmeyer, R. 
(eds) Die Fauuna Sudwest-Australiens. (G 
Fischer: Jena). 

HOOPER, J.N.A. 1996. Revision of the Microcionidae 
(Porifera: Poecilosclerida: Demospongiae), with 
description of Australian species. Memoirs of the 
Queensland Museum 40: 1-626. 

HOOPER, J.N.A. & BERGQLIIST P.R. 1 992. Cymbas- 
lella, a new genus of lamellate coral reef sponges. 
Memoirs ofthe Queensland Museum 32( 1 ): 99-1 37. 

HOOPER, J.N.A., LIST-ARMITAGE, S.E.. 
KENNEDY, J.A., COOK, S.D. & VALENTINE, 



C.A. 1999. Sponges of the Low isles. Great 
Barrier Reef: an important scientific site, or a case 
for mistaken identitv'? Memoirs of the Queensland 
Museum 44: 249-262. 

HOOPER, J.N.A. & WIEDENMAYER, E 1994. 
Porifera. Pp. 1-624. In Wells, A. (ed.) Zoological 
Catalogue of Australia. Vol. 12. (CSIRO Aus- 
tralia: Melbourne). 

KOBAYASHI, J., YUASA, K., KOBAYASHI, T, 
SASAKI, T & TSUDA, M. 1996. Jaspiferals A 
approx G, new cytotoxic isomalabaricane-type 
nortriierpenoids from Okinawan marine sponge 
Jaspisstellifera. Tetrahedron 52( 1 6): 5745-5750. 

LAIJBENFELS, M.W. de 1936. A discussion on the 
sponge fliuna of the Dry Tortugas in particular, 
and the West Indies in general, with material for a 
revision of the families and orders of Porifera. 
(Tortugas Laboratorv' Paper No. 467) Carnegie 
Institute of Washington 30: 1 -225. 
1950. The sponges of Kaneohe Bay, Oaliu. Pacific 

Science 4(1): 3-36. 
1954. The sponges of the West-Central Pacific. 

Oregon State Monographs, Zoology 7: 1-306. 
1957. New species and records of Hawaiian 
sponges. Pacific Science 11(2): 236-251. 

LENDENFELD, R. VON. 1896. Die Clavulina der 
Adria. Acta Academiee Csesarea; Leopoldino- 
Carolina: Gcmianicae Naturee Curiosorum 69: 1 -25 1 . 
1903. Tetraxonia. Pp. M68. In Schulze, F.E. (cd.) 
Das Tierrcich. Vol. 19. (Friedlander: Berlin). 

LEVI, C. 1973. Systematique de la classe de Demo- 
spongiaria (Demosponges). Pp. 577-631. In Brien, 
P,. Levi, C, Sara , M., Tuzet. O. & Vacelet, J. (eds) 
Traite de Zoologie, Anatomic, Systematique, 
Biologic. (Ser. ed. Grasse, P-P.) (Masson et Cie: 
Paris). 

McCaffrey, E.J. & ENDEAN, R. I985. Anti- 
microbial activit> of tropical and subtropical 
sponges. Marine Biology (Beriin) 89: 1-8. 

RAGONOT, E.L. 1887. Diagnoses of North American 
Phvcitidae and Galleriidae. (Paris). 

RAVI, B.N., WELLS, R.J. & CROFT, K.D. 1981. 
Malabaricane triterpenes from a Fijian collection 
of the sponge Jaspis stellifera. Journal of Organic 
Chemistry 46: 1998-2001. 

RAVI, B.N. & WELLS, R.J. 1982. Malabaricane 
triterpenes trom a Great Barrier Reef collection of 
the sponge 7as77w stellifera. Australian Journal of 
Chemistn 35: 39-50. ' 

RIDLEY, S.O. 1 884. Spongiida. Pp. 366-482, 582-635. 
In Report on the zoological collections made in 
the Indo-Pacific Ocean during the voyage of 
H.M.S. ^ Alert' 1881-2. (British Museum (Natural 
History): London). 

ROW, R.W.H. 1911. Reports on the marine biology of 
the Sudanese Red Sea, fi-om collections made by 
Cyril Crossland, M.A., B.Sc, F.Z.S. XIX. Report 
on the sponges collected by Cyril Crossland in 
1904-5. Part II. Non-Calc'area. Journal of the 
Linnean Society, Zoology 3 1 : 287-400. 



476 



MEMOIRS OF THE QUEENSLAND MUSEUM 



SANDERS, M., DIAZ, M.C. & CREWS, P. 1999. 
Taxonomic evaluation of jasplakinolide- 
containing sponges of the family Coppatiidae. 
Memoirs of the Queensland Museum 44: 
-525^532.. 

SCHMlDi; E.p. I862v Die Spoi^en des A<kiatischen 
Met£rBs.I*p, MS. (WilhelmEngelmann: Leipzig). 

li6S, Die SpDngien der Kiist von Algier. Mit 
Naditi^en zu den Spongien des Adriatlschen 
Meeies. (Drittes Supplement). Pp. 1-44. (Wil- 
helm Engelmann: Leipzig). 

1870. Grundziige einer Spongien-Fauna des 
Atlaritischen Gebietes. (Wilhelm Engelmann: 
Leipzig). 

SHAW, ME. 1927. On a collection of sponges from 
Maria Island, Tasmania. Proceediniis of the 
Zoological SocieW of London 1927: 41^9-430. 

SOEST, RAV.M. van '& BRAERMAM. J.C. 1999. 
Chemos> stemalicsofPorifera: are\ ievs. Memoirs 
of the Queensland Museum 44: 569-5S^K 

SOLLAS, WJ. 1 886. Preliminary account of the letrac- 
tinellid sponges dredged by H.M.S. 'Challenger', 
1872-1876. Part 1. The Chorislida. Scientific 
Proceedings, Royal Dublin Societ> 5: 177-199. 
1888. Report on the Tetractinellida collected by 
H.M.S. Thallenger\ during, the years 1873- 
1876* Pp. 1-458. In R6|m^ on the Sciendiic 
Results of tiie Voyage of MMS. ^Challenger', 
Zoology Vol. 25(63). 



THIELE, J. 1 899. Studien uber Pazifischen Spongier. 
Zoologica, Stuttgart 24(2): 1-33. 
1900. Kieselschwamme von Temate, 1. Abhhand- 
lungen der Senckenbeigtschen Naturforschenden 
Gesellseliaft^S: 19-80- 

1903. KieselschwilnimB von Ternate. IL 
Abhandiungen der Senokenbergischen 
Naturforschenden Gesellschait 25: 953^968. 

TOPSENT, E. 1898. Introduction 1 T^de mono- 
graphique des Monaxonides de France. 
Classification des Hadromerina. Archives de 
Zoologie Exp^rimentale et Generale ( 3 ifi : 9 1 - 1 1 3 . 

1904. Spongiaires des Azores. Resultais des 
Campagnes Scientitlques sur son Yacht par 
Albert ler Prince Souverain dc Monaco 25: 
1-280. 

I SUDA. M., ISHIBASHI, M, AGEMl, K., SASAKI, 
T. & KOBA^'ASHIJ. MWI . Stellitcrins A-E new 
antineoplastic isomalabancane triterpenes from 
ihe Okinawan marine sponge Jaspis steUyem* 
Tetrahedron 47(12-13): 2181-2194. 

WIEDENMAYER, F. 1989. Demospongiae (Porifera) 
from northern Bass Strait, southern Australia. 
Memoirs ofthe Museum of Victoria 50( 1 ): 1 -242. 

WILKINSON, C.R., SUMMONS, RJE, * EVANS. 
1999. Nitrogen fixation in symbiotic marine 
sponges: ecological sigmlicdnce and difQculttes 
in detectio^i. Memoirs ofthe QueenslandMtiSeum 
44; 6<57-673, 



PERCA LINEATA AND R VITTATA ESTABLISHED AS VALID SPECIES OF 
PLECTORHINCHUS (PERCIFORMES: HAEMULIDAE) 



JOHN E. RANDALL AND JEFFREY W. JOHNSON 

Randall, J.E. & Johnson, J.W. 2000 06 30: Perca lineata and P. vittata established as valid 
species of Plectorhinchus (Perci formes: Haemulidae). Memoirs of the Queensland Museum 
45(2): 477-482. Brisbane. ISSN 0079-8835. 

The Linnaean fishes Perca lineata and P. vittata are established as valid species of the 
haemulid genus Plectorhinchus. Plectorhinchus goldmanni (Bleeker) is a junior synonym of 
P. Uneatus, and P. orientalis (Bloch) is a junior synonym of P. vittatus. Mcristic data are 
tabulated to provide an additional basis for separating four other striped species of 
Plectorhinchus that at some stage resemble either P. lineatus or P. vittatus. □ Linnaeus, 
fishes, haemulidae, Plectorhinchus, goldmanni^ lineatus^ orientalis, vittatus. 

J.E. Randall, Bishop Museum, J 525 Bernice Street, Honolulu, Hawaii 96817-2704. USA; 
J. W. Johnson, Queensland Museum, PO Box 3300, South Brisbane 4 ] 1 , Australia: 2 1 April 
2000. 



Many fishes of the haemulid genus Plecto- 
rhinchus undergo remarkable changes in 
colouration with growth. There has been 
considerable confusion, particularly in the 
identification of some striped species. In the 
literature, juveniles have often been mismatched 
vvith adults, and various life stages have been 
incorrectly described as valid species, often with 
the aduh or Juvenile form unknown. In the latest 
review of the family, Smith (1962) failed to 
correctly associate many of the species with their 
synonyms, and he described G gaterinoides (a 
junior synonym ofP lineatus). He also presented 
a figure depicting seven colour phases of R 
orientalis from 135 to 550mm in length. The 
second to fourth specimens in the figure (fig. 1 5 
B, C and D) range from 1 75 to 233mm and clearly 
illustrate blotched individuals, howe\'er our studies 
indicate that specimens of this size are often, if 
not usually, striped as in larger specimens. The 
purpose of this paper is to validate two Linnaean 
species, both of which have a juvenile phase with 
horizontal stripes, and to compare meristic values 
of other similarly striped species of Plectorhinchus. 

Perca lineata and P. vittata are among the 29 
species classified in Perca by Linnaeus (1758). 
Both were described with reference to volumes 
of Museum Adophi Friderici as having five 
longitudinal white bands, but no locality details 
were provided. Cuvier in Cuvier and Valen- 
ciennes (1830: 309) was the first to correctly 
relate lineata with haemulid fishes when he 
placed it in the genus Diagranmra. He gave the 
dorsal-ray count for the species as XII, 20, thus 
resolving the count of XVII, 16 dorsal rays of 
Linnaeus by noting the small size and poor 



condition of the specimen and implying a 

miscount. Smith (1962: 495) suggested that the 
difference in dorsal-ray count might be a 
misprint. 

Fcm